Gayana Bot. 71(1):
68-107, 2014
71(1), 2014
ISSN 0016-5301
Revision of the genus Pseudognaphalium (Asteraceae, Gnaphalieae)
in Chile
Revisión del género Pseudognaphalium (Asteraceae, Gnaphalieae) en Chile
SUSANA E. FREIRE1, 2*, NÉSTOR D. BAYÓN2, CARLOS M. BAEZA3, DANIEL A. GIULIANO2 & CLAUDIA MONTI2
1
Instituto de Botánica Darwinion, Labardén 200, CC 22, B1642HYD San Isidro, Buenos Aires, Argentina.
Área de Botánica, Departamento de Ciencias Biológicas, Facultad de Ciencias Agrarias y Forestales, Universidad Nacional
de La Plata, Avda. 60 entre 116 y 118, 1900 La Plata, Argentina.
3
Facultad de Ciencias Naturales y Oceanográficas, Departamento de Botánica, Casilla 160-C, Universidad de Concepción,
Concepción, Chile.
*sfreire@darwin.edu.ar
2
ABSTRACT
The genus Pseudognaphalium is one of the largest genera of the tribe Gnaphalieae (Asteraceae) with about 90 species with
worldwide distribution. The vast majority of species are found in South, Central and North America, but some species also
occur in Asia and Africa. This work includes the revision of 12 species of Pseudognaphalium from Chile, P. aldunateoides,
P. cabrerae, P. cheiranthifolium, P. cymatoides, P. gayanum, P. lacteum, P. landbeckii, P. munoziae, P. psilophyllum, P.
remyanum, P. tarapacanum, and P. viravira. A key is provided for their identification, as well as morphological description,
illustration, geographic distribution and habitat, distribution maps, phenology, additional material examined, distinguishing
characters and taxonomic affinities for each taxon. The new combination Pseudognaphalium cabrerae (S.E. Freire) S.E.
Freire, N. Bayón, C. Baeza, Giuliano & C. Monti, is proposed. The following taxa are newly synonymized: Gnaphalium
diminutivum, Pseudognaphalium perpusillum, with P. aldunateoides; G. canum, G. ulophyllum, P. moelleri with P.
cymatoides; P. heterophyllum, P. heterotrichum, G. pseudohelichrysum, P. robustum with P. gayanum; P. glandulosum with
P. psilophyllum; G. longifolium, G. subnudum, P. andicola, P. coquimbense, P. illapelinum, P. montevidense, P. pratense,
with P. viravira. Lectotypes are newly designated for Gnaphalium acutifolium, G. araucanum, G. argyrolepis, G. canum,
G. cymatoides, G. glandulosum, G. heterophyllum, G. illapelinum, G. lacteum, G. landbeckii, G. moelleri, G. ramosum, G.
remyanum, G. subnudum, G. tarapacanum, G. ulophyllum, and G. viravira. A neotype is designated for G. psilophyllum.
KEYWORDS: Gnaphalium, lectotypification, neotype, South America, synonyms, taxonomy.
RESUMEN
El género Pseudognaphalium, con cerca de 90 especies de distribución cosmopolita, es uno de los géneros de la tribu
Gnaphalieae con mayor número de especies, principalmente distribuidas en América del Sur, Central y del Norte, aunque
algunas también están presentes en Asia y África. Este trabajo comprende la revisión de 12 especies de Pseudognaphalium
para Chile, P. aldunateoides, P. cabrerae, P. cheiranthifolium, P. cymatoides, P. gayanum, P. lacteum, P. landbeckii, P.
munoziae, P. psilophyllum, P. remyanum, P. tarapacanum y P. viravira. Se incluye una clave para identificarlas y para cada
una de las especies se brinda descripción, ilustración, fenología, distribución geográfica y hábitat, mapa de distribución,
material adicional examinado y afinidades taxonómicas. Sobre la base del estudio de los materiales tipo, se establecen los
siguientes nuevos sinónimos: Gnaphalium diminutivum, Pseudognaphalium perpusillum, con P. aldunateoides; G. canum,
G. ulophyllum, P. moelleri con P. cymatoides; P. heterophyllum, P. heterotrichum, G. pseudohelichrysum, P. robustum,
con P. gayanum; P. glandulosum con P. psilophyllum; G. longifolium, G. subnudum, P. andicola, P. coquimbense, P.
illapelinum, P. montevidense, P. pratense, con P. viravira; se designan lectotipos para las siguientes especies: Gnaphalium
acutifolium, G. araucanum, G. argyrolepis, G. canum, G. cymatoides, G. glandulosum, G. heterophyllum, G. illapelinum,
G. lacteum, G. landbeckii, G. moelleri, G. ramosum, G. remyanum, G. subnudum, G. tarapacanum, G. ulophyllum, G.
viravira y se designa el neotipo para G. psilophyllum.
PALABRAS CLAVE: Gnaphalium, lectotipificación, neotipo, sinónimos, Sud América, taxonomía.
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Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
INTRODUCTION
The cosmopolitan genus Pseudognaphalium Kirp. was
based on a Mexican species, Gnaphalium oxyphyllum
DC. It is one of the most species-rich genera of the tribe
Gnaphalieae (Asteraceae) with about 90 species of dwarf
prostrate to robust erect herbs. The vast majority of species
are distributed in South, Central and North America, but
some species also occur in Asia and Africa (Hilliard & Burtt
1981, Anderberg 1991, Bayer et al. 2007).
Pseudognaphalium was treated in regional works or
floral treatments, such as Rémy (1849, as Gnaphalium L.)
and Reiche (1905, as Gnaphalium) for Chile; Cabrera (1963,
1971, 1974, 1978, as Gnaphalium) for Argentina; Hilliard
(1983) and Hilliard & Burtt (1981) for South Africa; Dillon &
Sagástegui-Alva (1991a, b, as Gnaphalium) for Peru; Freire
(1998, as Gnaphalium) for Paraguay; Nesom (2001, 2004,
2006) and Ballard et al. ( 2004) for North America; Deble
& Marchiori (2006), and Freire et al. (2011) for Brazil; Hind
(2011) for Bolivia, and Chen et al. (2011) for Asia.
Cladistic analysis of morphological characters
(Anderberg 1991) and recent results from molecular studies
(e.g. Ward et al. 2009) have been interpreted as providing
support for recognizing Pseudognaphalium as a distinct
genus from Gnaphalium. The genus Pseudognaphalium
includes taxa with monochromous phyllaries with divided
stereomes, monomorphic pappus bristles, clusters of
capitula arranged in corymbs or panicles, and the achenes
either glabrous or with short oblong myxogenic duplex hairs.
In contrast, Gnaphalium includes taxa with polychromous
phyllaries with entire stereomes, dimorphic pappus bristles,
few capitula in sometimes leafy clusters, and pilose achenes
with oblong non-myxogenic duplex hairs (Drury 1970,
under ‘achyroclinoid’ cudweeds; Hilliard 1983, Hilliard
& Burtt 1981, Anderberg 1991). Furthermore, these and
other more recent molecular studies (Smissen et al. 2011,
Nie et al. 2013) noted that Pseudognaphalium is more
closely related to Helichrysum Mill. and Anaphalis DC.
than to Gnaphalium. Morphologically Pseudognaphalium,
including Laphangium (Hilliard & B.L. Burtt) Tzvelev,
resembles the genera Helichrysum and Anaphalis in having
divided stereome, receptacle smooth or honeycombed,
achenes glabrous or short pilose, and pappus uniseriate
with barbellate setae (Hilliard & Burtt 1981). It differs from
Helichrysum, however, in having pistillate florets always
outnumbering the bisexual ones (vs. pistillate florets not
outnumbering the bisexual florets), and from Anaphalis, in
having bisexual central florets (vs. functionally male).
Since the first taxonomic treatments of Chilean species
of Pseudognaphalium (under Gnaphalium, including
Gamochaeta Wedd.) were carried out by Rémy (1849),
Philippi (1895), and Reiche (1905), new species were
described or redescribed without a new broad taxonomic
evaluation or revision. Catalogues concerning aspects of
nomenclature of Pseudognaphalium (under Gnaphalium,
excluding Gamochaeta) by Marticorena & Quezada (1985),
and Freire & Iharlegui (2008) clarified synonyms and
superfluous names. Nevertheless, the revision of species in
Chile highlighted the need for further taxonomic adjustments.
The main goal of this study was to revise Chilean
species of Pseudognaphalium, considering morphological
characters and geographic distribution. The present study
also fully describes, illustrates, and provides a key to twelve
species occurring in Chile.
MATERIALS AND METHODS
The study is based on herbarium material from the following
herbaria (abbreviations according to Thiers 2011): BAB,
CONC, GH, LP, LPAG, MO, NY, PH, SGO, and SI.
Electronic images of type specimens at E, G-DC, L, P, SGO,
were also studied. Pappus bristles were observed after boiling
in water and staining with 2% safranin. Characteristics of
phyllaries and trichomes were observed and recorded in
cleared samples using Dizeo de Strittmatter’s technique
(1973). Observations were carried out using Gemalux
(Light Microscopy), equipped with a photographic camera
PAL CCD. Trichome terminology follows Ramayya (1962).
Leaf descriptions were made utilizing the terminology of
Harris & Harris (1994).
MORPHOLOGY AND MICROMORPHOLOGY
HABITAT AND STEMS. Most Chilean species of
Pseudognaphalium are perennial herbs or subshrubs,
more rarely biennials or annuals, i.e. P. aldunateoides,
P. cymatoides, ranging from few cm (P. aldunateoides,
P. lacteum) to up to 110 cm tall (P. gayanum), simple (or
2–3, e.g. P. cabrerae) or more usually with primary stem
branching at ground level only, and erect or ascending
unbranched stems from a decumbent base.
PUBESCENCE. Pubescence in Chilean species of
Peudognaphalium consists of two basic trichome types. The
type of pubescence is often useful alone or in combination
with other characters in distinguishing species (e.g., P.
cheiranthifolium from P. cabrerae). Two basic types of
hairs, present on stems, leaves, and phyllaries, are described
below.
1. Long uniseriate eseptate flagellate trichomes (Fig. 1A).
This type of trichome consists of one or two foot cells,
two-seven-celled stalks or stipes, and a single very long,
flagellate, eseptate apical cell. This trichome type is present
in all Chilean species of Pseudognaphalium, and especially
in Pseudognaphalium aldunateoides, P. cabrerae, P.lacteum,
P. landbeckii, P. remyanum, and P. viravira, where they
cover almost the entire surface of the leaf, with few short
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Gayana Bot. 71(1), 2014
glandular trichomes hidden under the wool.
2. Biseriate glandular trichomes (Fig. 1B, C). Comprised of
2 rows of cells in the body, with three to many cells in each
row, enclosed by a persistent or collapsed cuticular vesicle.
These trichomes vary from short clavate to long biseriate,
0.05–0.5 mm long. Glandular hairs are widespread in all
the taxa studied, and especially in Pseudognaphalium
cymatoides, P. gayanum, P. munoziae, P. psilophyllum,
and P. tarapacanum, where they cover almost the entire
surface of the leaf, with few flagellate trichomes. Only in P.
cheiranthifolium, long glandular trichomes are raised over
woolly trichomes.
ACHENIAL PUBESCENCE. Short pilose achenes occur in a few
Chilean species, i.e., P. aldunateoides, P. cymatoides, P.
gayanum (rarely P. tarapacanum, P. viravira). These are
twin hairs or duplex myxogenic hairs and consist of two
small, square basal cells and two short apical cells rounded
at the apex and equal in length (Fig. 2A, D). The remaining
taxa have glabrous achenes with smooth or papillose
epidermis (Fig. 2B, C, E).
LEAVES. All taxa analyzed have alternate, usually
approximated at the base, occasionally rosulate
(Pseudognaphalium munoziae), sessile leaves. The leaves
are always simple, predominantly linear, linear-lanceolate,
lanceolate or obovate. Only two species have conspicuously
undulate margins i.e. Pseudognaphalium cymatoides
and P. gayanum. In the remaining taxa the leaf margin is
predominantly flat. The leaf base is often stem-clasping and
decurrent.
INFLORESCENCE. The capitula in Chilean Pseudognaphalium
are arranged in a cymose pattern (Endress 2010). The
first-order axis (branch) has two short second-order axes
(branches), and these short lateral branches of each axis have
two or three branching orders, resulting a dense terminal
cluster (Fig. 3A, P. tarapacanum, P. viravira). However
in most species, clusters or cymose units are arranged in
thyrsoid inflorescence (Troll 1964-1969, Weberling 1985,
Rua 1999, Endress 2010) of three cymose units, commonly
called ‛corymbs’ (Fig. 3B, P. aldunateoides, P. cymatoides,
P. cheiranthifolium, P. cabrerae, P. gayanum, P. lacteum,
P. psilophyllum, P. viravira) or with more cymose units,
commonly called ‛panicle’ (Fig. 3C, P. aldunateoides,
P. cymatoides, P. gayanum, P. lacteum, P. landbeckii, P.
munoziae, P. psilophyllum, P. remyanum).
CAPITULA. The capitula of Pseudognaphalium are
heterogamous and discoid, containing in Chilean species
up to (51, 74) 95–212 florets with pistillate flowers
usually more than 10 times as many as bisexual flowers.
Campanulate involucres are characteristic of most species
of this study, ranging from 2 to 6 mm high.
PHYLLARIES. The phyllaries are arranged in 3 or 4 imbricate
series, with the inner equalling or slightly exceeding the
flowers, and not conspicuously longer than the outer.
The outer phyllaries are dorsally glandular pubescent and
the inner glandular or glabrous. The phyllary lamina is
monochromous opaque-white or hyaline, whitish, yellowish
or brownish and shiny, and the stereome is divided in all
taxa studied (Fig. 4A).
FIGURE 1. Foliar trichomes of Pseudognaphalium. A, Flagellate trichome (P. viravira, Teillier & Márquez 5315, CONC); B, Long biseriate
glandular trichomes (P. gayanum, Werdermann 470, SI); C, Short biseriate glandular trichome (P. gayanum, Werdermann 470, SI). Scale
bars: A−C= 0.1 mm.
FIGURA 1. Tricomas foliares de Pseudognaphalium. A, Tricoma flagelado (P. viravira Teillier & Márquez 5315, CONC); B, Tricoma
glandular biseriado largo (P. gayanum, Werdermann 470, SI); C, Tricoma glandular biseriado corto (P. gayanum, Werdermann 470, SI).
Escalas: A−C= 0,1 mm.
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Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
FIGURE 2. Achenes of Pseudognaphalium. A, Short pilose achene (P. aldunateoides, Urtubey et al. 700, SI); B, Achene with papillose
epidermis (P. psilophyllum, Werdermann 1100, SI). C, Surface view of imbricate achenial papillae (P. viravira, Matte 745, CONC). D,
Achene with short oblong duplex hairs (P. aldunateoides, Mihoc 6227, CONC). E, Glabrous achene with smooth epidermis and papillose
epidermis at the apex (P. munoziae, Moreira et al. 1676, SGO). Scale bars: A−E= 0.1 mm.
FIGURA 2. Aquenios de Pseudognaphalium. A, Aquenio cortamente piloso (P. aldunateoides, Urtubey et al. 700, SI); B, Aquenio con
epidermis papilosa (P. psilophyllum, Werdermann 1100, SI). C, Vista superficial de la epidermis del aquenio con papilas imbricadas (P.
viravira, Matte 745, CONC). D, Aquenio con pelos dobles cortos y oblongos (P. aldunateoides, Mihoc 6227, CONC). E, Aquenio glabro
con epidermis lisa, sólo papilosa en el ápice (P. munoziae, Moreira et al. 1676, SGO). Escalas: A−E= 0.1 mm.
FIGURE 3. Inflorescences of Pseudognaphalium. A, Capitula arranged in dense terminal clusters (P. tarapacanum, P. viravira); B, Capitula
in dense clusters arranged in corymbs (P. aldunateoides, P. cheiranthifolium, P. cymatoides, P. cabrerae, P. gayanum, P. lacteum, P.
landbeckii, P. psilophyllum, P. viravira); C, Capitula in dense clusters arranged in thyrsoid inflorescence (P. aldunateoides, P. cymatoides,
P. gayanum, P. lacteum, P. munoziae, P. psilophyllum, P. remyanum).
FIGURA 3. Inflorescencia de Pseudognaphalium. A, Capítulos agrupados en densos glomérulos terminales (P. tarapacanum, P. viravira);
B, Capítulos en densos glomérulos reunidos a su vez en cimas corimbiformes (P. aldunateoides, P. cheiranthifolium, P. cymatoides, P.
cabrerae, P. gayanum, P. lacteum, P. landbeckii, P. psilophyllum, P. viravira); C, Capítulos en densos glomérulos reunidos a su vez en
inflorescencias tirsoides (P. aldunateoides, P. cymatoides, P. gayanum, P. lacteum, P. munoziae, P. psilophyllum, P. remyanum).
71
Gayana Bot. 71(1), 2014
COROLLA. The corollas are whitish or yellowish. Corollas
of pistillate florets are filiform and corollas of bisexual
florets are tubular, scarsely broadened and glandulose
above, 5-lobed, usually whitish or yellowish (purplish in P.
aldunateoides).
ANTHERS. As well as most species of the tribe Gnaphalieae,
all the taxa of Pseudognaphalium in this study have an
apical ovate to lanceolate anther appendage and short basal
tails (Fig. 4B).
STYLE. As well as many taxa of the tribe Gnaphalieae
(e.g. Achyrocline, Gamochaeta), all the species of
Pseudognaphalium in this study have apically pilose styles,
with the two branches truncate to rounded at the apex (Fig.
4C).
PAPPUS. In all taxa studied pappus bristles are barbellate,
with apical cells rounded and inflated, and bases cohering
by patent cilia (Fig. 4D).
FIGURE 4. A, Phyllary showing the divided stereome (P. viravira, Barros 6076 LP); B, Anther (P. cheiranthifolium, Palma & Inostroza s.n.,
CONC); C, Style branch from bisexual floret (P. cheiranthifolium, Palma & Inostroza s.n., CONC); D, Pappus bristles (monomorphic)
showing tip cells and basal portion, left to right: pappus bristle of female, pappus bristle of bisexual floret (P. munoziae, Moreira et al. 1676,
SGO). Scale bars: A, B= 1 mm, C, D= 0.1 mm.
FIGURA 4. A, Filarios con esteroma dividido (P. viravira, Barros 6076 LP); B, Antera (P. cheiranthifolium, Palma & Inostroza s.n., CONC);
C, Ramas del estilo de una flor perfecta (P. cheiranthifolium, Palma & Inostroza s.n., CONC); D, Pelos del papus (monomórfico) mostrando
las células apicales y las cilias basales; a la izquierda: pelo del papus de una flor pistilada; a la derecha: pelo del papus de una flor perfecta
(P. munoziae, Moreira et al. 1676, SGO). Escalas: A, B= 1 mm, C, D= 0.1 mm.
TAXONOMIC TREATMENT
Twelve species of Pseudognaphalium occur in Chile,
representing about 15% of the known species for the genus.
Pseudognaphalium Kirp., Trudy Bot. Inst. Akad. Nauk
S.S.S.R., Ser 1, Fl. Sist. Vyssh. Rast. 9: 33. 1950. TYPE:
Gnaphalium oxyphyllum DC. = Pseudognaphalium
oxyphyllum (DC.) Kirp.
Ser. 1, Fl. Sist. Vyssh. Rast. 9: 33. 1950. TYPE: Gnaphalium
heterotrichum Phil. = Hypelichrysum heterotrichum (Phil.)
Kirp. = Pseudognaphalium heterotrichum (Phil.) Anderb.
Gnaphalium L. sect. Calolepis Kirp., Bot. Mater. Gerb.
Bot. Inst. Komarova Akad. Nauk S.S.S.R. 20: 309. 1960.
Pseudognaphalium subgen. Laphangium Hilliard & B.L.
Burtt, Bot. J. Linn. Soc. 82(3): 205. 1981. Laphangium
(Hilliard & B.L. Burtt) Tzvelev, Byull. Moskovsk. Obshch.
Isp. Prir. Otd. Biol. 98(6): 105. 1993[1994].
Gnaphalium L., Sp. Pl. 2: 850. 1753, p.p. excl. type.
Hypelichrysum Kirp., Trudy Bot. Inst. Akad. Nauk SSSR,
72
Gnaphalium Kirp. subgen. Laphangium (Hilliard &
B.L. Burtt) P.D. Sell, Fl. Gr. Brit. Ireland 4: 555. 2006.
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
TYPE: Gnaphalium luteoalbum L. = Pseudognaphalium
luteoalbum (L.) Hilliard & B.L. Burtt.
Annual, biennial or perennial herbs, woolly or glandularwoolly. Leaves alternate, entire, sessile, often stem-clasping
and decurrent. Capitula small, heterogamous, discoid, sessile
or short pedunculate, in small clusters arranged in corymbs
or panicles. Involucre often campanulate; phyllaries in
3–4-series, papery, phyllary lamina monochromous opaquewhite or hyaline and shiny, stereome divided. Receptacle
smooth or honeycombed. Florets many, pistillate florets
outnumbering the bisexual; pistillate corollas filiform
or narrowly tubular; bisexual corollas tubular, scarsely
broaded above, 5-lobed, all corollas yellowish or whitish,
lobes glandular and often yellowish or whitish. Anthers
with a small obtuse apical anther appendage; tails slightly
longer or shorter than the filament collar. Style branches
truncate and penicillate. Achenes glabrous often with
imbricate microcopic papillae or pilose with short duplex
myxogenic hairs. Pappus bristles monomorphic, barbellate,
apical cells sometimes inflated, bases cohering by patent
cilia. Chromosome number: 2n = 14 (Turner, 1970; Mehra
& Remanandan, 1975; Keil, 1981; Groves, 1977); 2n = 28
(Keil & Stuessy, 1977).
Worldwide distribution, mostly American, some African,
and Asian species. About 90 species of which twelve grow
in Chile between sea level and 4500 m.
KEY TO THE SPECIES OF PSEUDOGNAPHALIUM IN CHILE
1. Dwarf herbs; stems prostrate, branched. …………..………….....................................................................................………… P. lacteum
1’. Dwarf or large herbs; stems erect or ascending, branched or unbranched.
2. Basal leaves rosulate, longer than the upper leaves; upper leaves alternate. Stems 4–6(15) cm tall. .....................................P. munoziae
2’. Basal and upper leaves alternate.
3. Leaf blade margins conspicuously undulate.
4. Weak stems 7–50 cm tall, branched; involucre 2–3 mm high……….......………………....................………….P. cymatoides
4´. Robust stems 30–110 cm tall, unbranched; involucre 4–5 mm high……………………..................………..……P. gayanum
3’. Leaf blade margins flat (occasionally some with undulate margins).
5. Leaves glandular or arachnoid-glandular.
6. Upper leaves broad at base…………….......................................................................................….………P. tarapacanum
6’. Upper leaves not broad at base.............…...............................................................................…………… P. psilophyllum
5´. Leaves usually whitish-woolly with short glandular trichomes hidden under the woolly trichomes (rarely with long glandular
trichomes raised over woolly trichomes).
7. Dwarf plants (stems 1–10 cm). Capitula 3-5(7) arranged in corymbs or narrow panicles. Achenes short pilose ………
…….....................................................................................................................................................… P. aldunateoides
7’. Usually large plants; stems > 10 cm tall (when less than 10 cm tall, basal leaves approximate). Capitula numerous.
Achenes glabrous or short pilose.
8. Caulinar leaves oblong, apex obtuse…...............................................................................................…P. landbeckii
8’. Caulinar leaves linear, linear-lanceolate, lanceolate or linear-obovate, apex acute.
9. Leaves linear-lanceolate or lanceolate.
10. Stem and leaves whitish, woolly with long glandular trichomes raised over the woolly trichomes. Leaves
concolorous…..…………………….......................................................................….P. cheiranthifolium
10’. Stems and leaves whitish, woolly, with short glandular trichomes hidden under the woolly trichomes.
Leaves slightly discolorous to discolorous………...........................................................……P. cabrerae
9´. Leaves linear or linear-obovate.
11. Stem branched above the base. Leaves arachnoid, adaxial surface green. Capitula arranged in corymbs
or lax panicles. ….........................................................................................................…P. remyanum
11´. Multistemmed plants with unbranched stems. Leaves usually whitish woolly. Capitula usually
arranged in dense terminal clusters.....…............................................................................P. viravira
1. Pseudognaphalium aldunateoides (J. Rémy) C. Monti,
N. Bayón & S.E. Freire, J. Bot. Res. Inst. Texas 7(1): 196.
2013. Gnaphalium aldunateoides J. Rémy, in Gay Fl. Chil.
4(2): 232. 1849. TYPE: Chile. ‘Se cría en los cerros de
Santiago y florece en setiembre [Gay]’ (P). Fig. 5.
Gnaphalium insulare Phil., Bot. Zeitung (Berlin) 14: 645.
1856. TYPE: CHILE. Islas de Juan Fernández, Más Afuera,
X-1854, Germain s.n. (holotype, SGO 64416!; isotypes, B,
probably destroyed, = photo F15103!, LP 001926 fragment
ex SGO!).
Gnaphalium perpusillum Phil., Linnaea 29: 6. 1858, syn.
nov. Pseudognaphalium perpusillum (Phil.) C. Monti, N.
Bayón & S.E. Freire, J. Bot. Res. Inst. Texas 7(1): 197.
2013. TYPE: CHILE. Coquimbo: Sotaqui, Jan 1837, C. Gay
739 (holotype, SGO 64448!).
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Gayana Bot. 71(1), 2014
FIGURE 5. Pseudognaphalium aldunateoides. A, A’, Habits; B, Leaf pubescence; C, D, Foliar trichomes; E, Capitulum; F, Phyllaries; G,
Pistillate floret; H, Bisexual floret. I, Distribution [A, C–E, H, Urtubey et al. 700, SI; A’, B, Boelcke 2414, LP; F, Ricardi 5445, CONC].
FIGURA 5. Pseudognaphalium aldunateoides. A, A’, Plantas; B, Pubescencia de la hoja; C, D, Tricomas foliares; E, Capítulo; F, Filarios;
G, Flor pistilada; H, Flor perfecta. I, Distribución [A, C–E, H, Urtubey et al. 700, SI; A’, B, Boelcke 2414, LP; F, Ricardi 5445, CONC].
74
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
Gnaphalium phaeolepis Phil., Linnaea 33: 168. 1864.
TYPE: Chile. without locality, year 1865, Frumau 63
(lectotype designated by Monti et al. (2013: 196), SGO
64452!; isolectotypes, B (probably destroyed), = photo
F15112!, SGO 44946!, 44947!).
Gnaphalium diminutivum Phil., Linnaea 33: 167. 1864.
syn. nov. TYPE: Chile. Colchagua, Cerro del Volcán
[ad originem fluvii Tinguiguirica], II-1861, W. Diaz s.n.
(lectotype designated by Monti et al. (2013: 197), SGO
64454!; isolectotype SGO 44923!).
Dwarf annual herbs, 1–10 cm tall, multistemmed (single
stem in very small plants), stems erect or ascending, usually
unbranched, lanuginose. Stem leaves 6–16 × 1–3(4) mm,
linear or linear-oblong, margin flat, apex acute or subobtuse,
mucronate; basal leaves 8–9 × 2.5–3 mm; concolorous,
whitish-woolly on both faces, trichomes of 2 types, long
eglandular eseptate, 3–8-celular, and few short glandular
biseriate trichomes, hidden under the wool. Capitula few,
3–5(7) in clusters arranged in corymbs or narrow panicles,
clusters subtended by linear leaves. Involucre campanulate,
3–3.2 × 2.5–3 mm. Phyllaries 3–4-seriate, lamina opaque
white or hyaline whitish at apex; outer phyllaries 1.8–2.8
× 0.6–1 mm, ovate, apex obtuse to acute; inner phyllaries
1.5–3 × 0.4–0.9 mm, linear-obovate, apex acute. Pistillate
florets 96–150, corolla white-yellowish, purplish at apex,
filiform, 0.9–1.7 mm long. Bisexual florets 4–13, corolla
white-yellowish, purplish at apex, tubular, 1–1.7 mm long.
Achenes 0.3–0.6 mm long, pilose, with short oblong duplex
hairs. Pappus bristles 1–1.8 mm long.
PHENOLOGY. Flowering and fruiting from October to March.
DISTRIBUTION AND HABITAT. Central and South Chile
(Coquimbo, Valparaíso, Metropolitana, Libertador General
Bernardo O´Higgins, Archipiélago Juan Fernández, Maule,
Biobío, Araucanía) and West Argentina (Chubut, Mendoza,
Neuquén, Río Negro, San Juan), growing in mossy places
and river margins, from sea level to 3700 m.
RELATIONSHIPS. Pseudognaphalium aldunateoides is easily
distinguished by its dwarf habit with erect to ascending
unbranched stems, linear leaves, few capitula in clusters
arranged in corymbs or narrow panicles, and short pilose
achenes.
OBSERVATIONS.
1. According to the protologue, Gnaphalium aldunateoides
was based on the specimen. ‘Se cría en los cerros de Santiago
y florece en setiembre’. We found at P (00704557, = photo!)
a collection, ‘Stiago. (Chili), Sbre. 1839, C. Gay 936’ which
is in accordance with the protologue (although no collection
number is mentioned there) and has the annotation ‘Type’.
Since we were able to see only one sheet, we prefer not to
select lectotype in this case.
2. Klatt (Linnaea 42: 121. 1878) mentioned Gnaphalium
rivulare, as a synonym of G. insulare Phil. According
to Philippi (Anales Univ. Chile 90: 18. 1895) it is a
misinterpretation by Klatt, who express that he found at
Berlin herbarium two specimens collected at Masafuera
island by Philippi, one named G. rivulare and the other one
G. insulare. Philippi (l.c.) stated that Klatt probably read
‛riv’ instead of ‛ins.’
3. Gnaphalium perpusillum Phil. was described on the
basis of its dwarf plants ca. 1.5 cm tall with single stem.
However, this is a variable feature throughout the range of
G. aldunateoides.
4. Gnaphalium diminutivum Phil. which was previously
placed into synonymy of G. perpusillum Phil. (Monti et al.
2013) is here considered as synonym of Pseudognaphalium
aldunateoides.
ADDITIONAL MATERIAL EXAMINED. CHILE. Región de
Coquimbo. Prov. Elqui, Paihuano, III-1946, Roesner 6508
(CONC); Prov. Limarí, Ovalle, Tulahuén, 14-I-1942, Barros
2258 (LP). Región de Valparaíso. Prov. Los Andes, Comuna
de Los Andes, cajón del río Colorado, vega de Lagunillas,
32°36’S, 70°20’W, 20-II-2002, Teillier 51112 (CONC);
Potrero Escondido, 22-II-1947, 3500 m, Boelcke 2424 dupl.
(LP); Prov. San Felipe de Aconcagua, Putaendo, Laguna
de Copín, 2620 m, 26-I-1955, Parra 37 (CONC); Prov.
Valparaíso, año 1891, Cuming s.n. (LP), Lago Peñuelas,
7-II-1968, Ricardi 5445 (CONC), Placilla, 33°07’S,
71°35’W, 28-XII-1993, Teillier 3224 (CONC). Región
Metropolitana de Santiago. Prov. Cordillera, San José del
Maipo, Cajón del Morales, entre la laguna y el glaciar, 27I-2002, Teillier & Márquez 5316 (CONC); Prov. Melipilla,
Alhué, Monte Cantillana, 2-I-1939, Barros 2272 (LP); Prov.
Santiago, vega entre Casa de Piedra Carvajal y Cerro La
Paloma, sector alto del estero de la Yerba Loca, 33°13’04’’S,
70°16’18.2’’W, 14-II-1999, Arroyo & Humaña 991714
(CONC); Bajando de La Parva, 2859 m, 24-I-2012, Urtubey
et al. 700 (SI). Región del Maule. Prov. Cauquenes, ca. 2 km
south of Chanco on west side of road, 35°45’S, 72°32’W, 50
m, 22-XI-1995, Bliss 2272 (CONC); Prov. Linares, Quella
Sur, 36°05’S, 72°08’W, 21-I-1991, Matthei & Quezada
1420 (CONC); Prov. Talca, Curepto, Puninque, 21-I-1926,
Barros 34 (LP); Curepto, 10 m, I-1925, Barros 649 (CONC);
Cordillera de Talca, El Picazo, 26-I-1939, Barros 2290 (LP).
Región del Biobío. Prov. Arauco, Contulmo, cerros orillas
L. Lanalhue, 30-XII-1949, Ricardi 9259 (CONC); Prov.
Biobío, Nacimiento, 26-XII-1930, Barros 45 (LP); Negrete,
a orillas del Bío-Bío, en la ribera norte del río, 19-II-1951,
Pfister 10502 (CONC); Prov. Concepción, Hualpén, 9-III-
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Gayana Bot. 71(1), 2014
1940, Junge 6767 (CONC); Prov. Ñuble, Ninhue, Río
Ninhue, km 32 desde San Gregorio a Pocillas, 190 m,
36°08’S, 72°05’W, 4-II-2003, Mihoc et al. 6227 (CONC).
Región de la Araucanía. Prov. Cautín, Metrenco, 85 m, XI1946, Gunckel 46993 (CONC); Temuco, Río Cautín, 1-XII1957, Montero 5728 (CONC, LP).
2. Pseudognaphalium cabrerae (S.E. Freire) S.E. Freire, N.
Bayón, C. Baeza, Giuliano & C. Monti, comb. nov.
Basyonym: Gnaphalium cabrerae S.E. Freire, Flora
Fanerogámica Argentina 14, Errata et corrigenda 4. 2000.
Gnaphalium fastigiatum Phil., Anales Univ. Chile 90:
15. 1895, nom. illeg., non Thunb., 1800 nec Schrank,
1824. Gnaphalium philippii Cabrera, Revista Mus. La
Plata, Secc. Bot. 4: 164. 194, nom. illeg., non Gandoger,
1918. Gnaphalium cabrerae S.E. Freire, Monogr. Syst.
Bot. Missouri Bot. Gard. 74(2): 1245. 1999, nom. inval.
Pseudognaphalium cabrerae (Freire) Deble, Balduinia 6:
29. 2006, comb. inval. Pseudognaphalium fastigiatum N.
Bayón, Bol. Soc. Argent. Bot. 48 (3-4): 599. 2013, nom.
illeg. TYPE: Chile. Santiago, Quinta Normal, III-1878,
R.A. Philippi s.n. (holotype, SGO 64449!; isotype, LP
001914!).). Fig. 6.
Annual herbs, 40–60 cm tall, stem usually solitary, erect,
rarely branched at the upper part, whitish-woolly. Leaves
35–70 × 4–7 mm, lanceolate, margin flat, apex longattenuate, acute, base decurrent 7-8 mm long; slightly
discolorous to discolorous, with upper surface lanuginose
and lower surface densely tomentose, trichomes of 2 types,
long eglandular eseptate, 3-4-celular, and few short biseriate
glandular trichomes hidden under the wool. Capitula
numerous in clusters arranged in corymbs. Involucre broadly
campanulate, 3–4 × 3–4 mm. Phyllaries 4-seriate, lamina
straw-coloured to brownish, hyaline and shiny, apex obtuse;
outer phyllaries 3–3.2 × 1.5–1.6 mm, ovate; inner phyllaries
3–3.3 × 0.5–0.8 mm, oblong-obovate. Pistillate florets ca.
198, corolla yellow, filiform, 3.2 mm long. Bisexual florets
ca. 11, corolla yellow, tubular, ca. 3.2 mm long. Achenes ca.
0.5 mm long, glabrous, epidermis papillose. Pappus bristles
ca. 2.8 mm long.
PHENOLOGY. Flowering from November to February.
DISTRIBUTION AND HABITAT. Central Argentina (Buenos Aires,
Catamarca, Córdoba, La Rioja, Mendoza, San Juan, San
Luis) and North, Central Chile (Tarapacá, Metropolitana,
Maule, Araucanía), growing from sea level to 2000 m.
RELATIONSHIPS. Pseudognaphalium cabrerae is closely
related to P. cheiranthifolium. Both species are woolly plants
76
with usually solitary stems, linear-lanceolate or lanceolate
leaf blades and numerous capitula in clusters arranged in
corymbs. Pseudognaphalium cabrerae can be distinguished
by its woolly leaves with short glandular trichomes hidden
under the wool (vs. woolly leaves with long glandular
trichomes raised over the wool in P. cheiranthifolium).
OBSERVATION
Recently, Bayón & Giuliano (2013) proposed the name
Pseudognaphalium fastigiatum N. Bayón as the correct
name for Gnaphalium fastigiatum Phil. (hom. illeg.) when
transferred to the genus Pseudognaphalium. However,
according to ICN (McNeill et al. 2012), Art.11.4., the
correct name is P. cabrerae, since the earliest legitimate
name of the taxon is Gnaphalium cabrerae S.E. Freire,
Flora Fanerogámica Argentina 14, Errata et Corrigenda 4.
2000. Since the name Pseudognaphalium cabrerae (S.E.
Freire) Deble is not validly published because it was based
on Gnaphalium cabrerae S.E. Freire, Monogr. Syst. Bot.
Missouri Bot. Gard. 74(2): 1245. 1999 (nom. inval.), we
propose the new combination Pseudognaphalium cabrerae
(S.E. Freire) S.E. Freire, N. Bayón, C. Baeza, Giuliano &
C. Monti, based on Gnaphalium cabrerae S.E. Freire, Flora
Fanerogámica Argentina 14, Errata et Corrigenda 4. 2000.
ADDITIONAL MATERIAL EXAMINED. CHILE. Región de Tarapacá.
Prov. El Tamarugal, Termas de Mamiña, 15-IX-1981, 2900
m, Montero 12045 (CONC). Región Metropolitana de
Santiago. Cerro Manquehue (Canal Lo Curro), 7-XII-1976,
Elgueta 546 (CONC). Región del Maule. Prov. Curicó,
Potrero Grande, 4-I-1927, Barros 1940 (CONC). Región
de la Araucanía. Prov. Malleco, Lonquimay, 20-II-1936,
Atanasio 706 (CONC).
3. Pseudognaphalium cheiranthifolium (Lam.) Hilliard &
B.L. Burtt, Bot. J. Linn. Soc. 82(3): 205. 1981. Gnaphalium
cheiranthifolium Lam., Encycl. [J. Lamarck & al.] 2(2):
752. 1788. TYPE: [Uruguay]. ‘Commerson à trouvé cette
plante à Monte-Video’ (P-Lam.?). Fig. 7.
Gnaphalium citrinum Hook. & Arn., Bot. Beechey Voy.: 31.
1830. TYPE: Chile. ‛Hab. Concepción’ (holotype, probably
K; isotype, SGO 72232!).
Gnaphalium acutifolium Phil., Anales Univ. Chile 90: 12.
1895. TYPE: Chile. Arauco. Curanilahue I-1893, Philippi
s.n. (lectotype, here designated, SGO 64401!; isolectotypes,
SGO 35992!, 44968!).
Gnaphalium araucanum Phil., Anales Univ. Chile 43:
502. 1873. TYPE: Chile. Biobío: Lebu, G. Volkmann s.n.
(lectotype, here designated, SGO 71287!; isolectotype,
SGO 71281!).
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
FIGURE 6. Pseudognaphalium cabrerae. A, Habit; B, Leaf pubescence; C, D, Foliar trichomes; E, Capitulum; F, Phyllaries; G, Pistillate
floret; H, Bisexual floret. I, Distribution [A, E, G, H, Barlett 20018, SI; C, D, Cabrera 187, LP; B, F, Cabrera 5189, LP].
FIGURA 6. Pseudognaphalium cabrerae. A, Planta; B, Pubescencia de la hoja; C, D, Tricomas foliares; E, Capítulo; F, Filarios; G, Flor
pistilada; H, Flor perfecta. I, Distribución [A, E, G, H, Barlett 20018, SI; C, D, Cabrera 187, LP; B, F, Cabrera 5189, LP].
77
Gayana Bot. 71(1), 2014
FIGURE 7. Pseudognaphalium cheiranthifolium. A, Habit; B, Leaf pubescence; C, D, Foliar trichomes; E, Capitulum; F, Phyllaries. G,
Distribution [A, E, from Cabrera 1978; B–D, Palma & Inostroza s.n., CONC; F, Finot & Leppe 97, CONC].
FIGURA 7. Pseudognaphalium cheiranthifolium. A, Planta; B, Pubescencia de la hoja; C, D, Tricomas foliares; E, Capítulo; F, Filarios; G,
Distribución [A, E, tomado de Cabrera 1978; B–D, Palma & Inostroza s.n., CONC; F, Finot & Leppe 97, CONC].
78
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
Gnaphalium paniculatum Bertero ex Colla, Mem.
Reale Accad. Sci. Torino 38: 17. 1835. Gnaphalium
cheiranthifolium Lam. var. paniculatum (Bertero ex Colla)
Skottsb., Kongl. Svenska Vetenskapsakad. Handl. 51(9):
5. 1914. TYPE: [Chile] ‘Cultum in H. Ripul. e seminibus
missis a BERTERO sine nomine specifico et lectis in
saxosis apricis Valparaiso, servatur hyeme in frigidario, et
floret secundo anno, ac raro diutius perdurat.’ (holotype,
probably TO, not seen).
Gnaphalium valdivianum Phil., Linnaea 29: 6. 1858. TYPE:
Chile. Valdivia, IV-1852, s. leg. (holotype, SGO 64374!).
Gnaphalium cheiranthifolium Lam. var. multiflorum J.
Koster, Blumea 5(3): 655. 1945. TYPE: Bolivia. Samaipata,
2000-2200 m, III-1911, T. Herzog 1758 (holotype, L, =
phot!; isotypes, B 10 0093571, = phot!, LP 001906!).
Perennial herbs, 40–80 cm tall, stem solitary (2 or 3),
erect, unbranched or branched at the upper part, woollyglandulose. Stem leaves 30–45 × 2–5 mm, lanceolate or
linear-lanceolate, margin flat, apex long-attenuate, acute,
base decurrent 6–13 mm long; basal leaves 40–80 × 3–8
mm, oblanceolate-spathulate, apex obtuse; concolorous,
adaxial surface glandulose-tomentose, trichomes of 2 types,
long eglandular eseptate, 3-celular, and long glandular
biseriate trichomes raised over the wool, abaxial surface
tomentose. Capitula numerous in clusters arranged in
corymbs. Involucre broadly campanulate, 4–4.5 × 5–6 mm.
Phyllaries 4-seriate, lamina yellow-greenish, hyaline and
shiny; outer phyllaries 3.8–4 × 1.7–1.8 mm, ovate, apex
subobtuse; inner phyllaries 4.3–4.5 × 0.6–1 mm, oblongobovate, apex acute. Pistillate florets 78–80, corolla yellow,
filiform, ca. 3 mm long. Bisexual florets 10–15, corolla
yellow, tubulose, ca. 3 mm long. Achenes 0.7–0.8 mm long,
glabrous, epidermis smooth or papillose. Pappus bristles ca.
3 mm long.
PHENOLOGY. Flowering from October to April.
VERNACULAR NAME. ‛Té de burro’.
DISTRIBUTION AND HABITAT. Southern South America, i.e.
southern Brazil (Paraná, Santa Catarina; Freire et al. 2011),
Bolivia (Hind 2011), Uruguay (Rocha, San José), Argentina
(Buenos Aires, Córdoba, Entre Ríos, Jujuy, La Pampa,
La Rioja, Salta, San Luis), and Chile (Arica y Parinacota,
Coquimbo, Valparaíso, Metropolitana, Libertador General
Bernardo O´Higgins, Maule, Biobío, Araucanía, Los Lagos,
Los Ríos, Archipiélago Juan Fernández). Commonly
collected on rocky and sandy soils, from sea level to 2500 m.
RELATIONSHIPS. Pseudognaphalium cheiranthifolium is
closely related to P. cabrerae. Both species are plants
with usually solitary stems, linear-lanceolate or lanceolate
leaf blades and numerous capitula in clusters arranged in
corymbs. Pseudognaphalium cheiranthifolium can be
distinguished by its concolorous woolly leaves with long
glandular trichomes raised over the wool (vs. slightly
discolorous to discolorous leaves, lanuginose on adaxial
surface and densely tomentose on abaxial surface, with short
glandular trichomes hidden under the wool in P. cabrerae).
OBSERVATIONS
1. According to the protologue, Gnaphalium cheiranthifolium
Lam., was based on the specimen ‘Commerson à trouvé
cette plante à Monte-Video’. We found at P the collection
Commerson 103, P 00704563, which is in accordance with
the protologue (although no collection number is mentioned
there). However, since we were unable to locate this
specimen at Lamarck’s herbarium, we prefer not to select
lectotype in this case.
2. According to the protologue, Gnaphalium araucanum
Phil., was based on the specimen ‘don Jerman Volkmann
halló esta especie cerca de Lebu, en la Araucania’. Muñoz
Pizarro (1960) cited SGO 64381 as the holotype. However,
only two other sheets ‘Lebu, Volkmann s.n.’ were located at
SGO, SGO 71281 and SGO 71287. We selected as lectotype
the specimen SGO 71287, which is better preserved.
3. According to the protologue, Gnaphalium paniculatum
Bertero ex Colla, was based on the specimen ‘Cultum in
H. Ripul. e seminibus missis a BERTERO sine nomine
specifico et lectis in saxosis apricis Valparaiso, servatur
hyeme in frigidario, et floret secundo anno, ac raro
diutius perdurat.’ We located three sheets, P 00704559, P
00704560, P 00704561, and one at LP 001936! ex P of the
specimen ‛Valparaiso, XII-1829, Bertero 1033’, which are
in accordance with the protologue (although no collection
number and date are mentioned there). Since we were
unable to see the Colla’s original material probably kept at
TO, we prefer not to select lectotype in this case.
ADDITIONAL MATERIAL EXAMINED. CHILE. Región de Arica
y Parinacota. Prov. Parinacota, camino entre Putre y
Socoroma, km 8, 3600 m, 18º12’S, 69º35’W, 21-III-1987,
Matthei & Rodríguez 299 (CONC). Región de Coquimbo.
Prov. Choapa, Carretera Panamericana, 7 km N de Los
Vilos, cerca de Agua Amarilla, 15 m, 31º51’S, 71º30’W,
2-XI-1974, C. Marticorena et al. 353 (CONC); Pichidangui,
30 m, 19-II-1966, Mahu 10335 (LP); Prov. Elqui, Quebrada
Las Trancas, 25 km S of Vicuña, 1350 m, 16-XI-1940,
Wagenknecht s.n. (MO 1294135); Prov. Limari, 1 km W
of road to Parque Nacional Fray Jorge, 30º37´S-71º33´W,
200 m, 4-XI-2006, Tepe et al. 1727 (CONC); Región de
Valparaíso. Prov. San Antonio, Rocas de Santo Domingo, 25
m, 20-X-1950, Pfister & Ricardi s.n. (CONC 9760); Prov.
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Gayana Bot. 71(1), 2014
Petorca, Carretera Panamericana, 3 km antes del puente
Guaquén, 15-X-1971, C. Marticorena et al. 1307 (BAB, LP,
CONC); Longotoma, 6-I-1970, Crisci 349 (LP); Papudo,
above coastal cliffs, 10-XI-2006, 32º30’21’’S, 71º28’08’’W,
Tepe et al. 1923 (CONC); Prov. Valparaíso, Cerros de L.
Verde, 11-II-1931, Garaventa 1965 (CONC); Valparaíso,
JBN de Viña del Mar, Quebrada Los Colectivos, 21-I-2005,
Novoa s.n. (CONC 162506); ca. 6 km N of Viña del Mar
on coast road, 70-100 m, 33º00’S, 71º35’W, 24-XII-1978,
J. & A. Solomon 4226 (MO); Quintero, 20 m, XI-1952,
Hochleitner s.n. (CONC 116164); ídem, 20 m, IX-1923,
Werdermann 37 (SI); Quintero, 6-II-1952, Levi Heins 154
(CONC); Quintero, Los Molles, 32º46’S, 71º32’W, 20 m,
XII-1951, Gunckel 23443 (CONC); ídem, 20 m, Gunckel
40589 (CONC); Ritoque, Loma de las Piedras, 32º46’S,
71º32’W, 10 m, II-1963, Gunckel 40215 (CONC). Región
Metropolitana de Santiago. Prov. Santiago, San Antonio, 10
m, 16-X-1927, Montero 222 (CONC). Región del Libertador
General Bernardo O´Higgins. Prov. Cardenal Caro, Tanumé,
Potrero Ramírez, 28-I-2005, 34º12’S, 71º57’W, Vera & Ortiz
21 (CONC); Tanumé, Los Romeros, 34º12´S-71º57´W, 350
m, 28-I-2005, Vera & Ortiz 151 (CONC); Loma El Camino,
350 m, 30-I-2005, 34º12’S, 71º57’W, Vera & Ortiz 136
(CONC). Región del Maule. Prov. Cauquenes, Reserva
Nacional Los Queules, 462 m, 35º59’16’’S, 72º41’35’’W,
6-I-2005, A. Marticorena & Jiménez 214 (CONC); Prov.
Curicó, Llico, 15-XII-1938, Barros 2279 (LP); 8 km al N de
Reserva Nacional Laguna Torca, quebrada y cerros frente
a la playa, 34º42´S-72º02´W, 20 m, 13-XII-2000, Finot &
Leppe 97 (CONC). Prov. Talca, Constitución, Los Molinos,
30 m, 25-X-1958, Barnier 225 (CONC). Región del Biobío.
Prov. Arauco, camino a Playa Quiala, 212 m, 37º20’S,
73º33’W, 13-I-2001, Parra & Torres 223 (CONC); 2,5 km
al N de Quidico, 45 m, 38º12’S, 73º27’W, 21-II-2001, Parra
& Torres 819 (CONC); camino a Yani, 179 m, 37º21’S,
73º37’W, 13-I-2001, Parra & Torres 163 (CONC); a 13 km
del cruce a Ranquilco, en camino Lebu-Los Álamos, 150
m, 37º44’S, 73º30’W, 23-II-2001, Parra & Torres 1026
(CONC); Caleta Rumena, 8 m, 37º10’S, 73º36’W, 3-I2001, Baeza & López 2759 (CONC); Prov. Biobío, camino
de Biobío a Santa Bárbara, cerca del Puente Río Queuco,
450 m, 37º50’S, 71º40W, 14-III-1976, C. Marticorena et
al. 1006 (CONC); Prov. Concepción, Rocoto, La Puntilla,
30 m, 22-X-1970, Palma & Inostroza s.n. (CONC 35172);
Rocoto, 30 m, 15-XI-1967, Villarroel & Weldt 123 (CONC);
Concepción, 10 m, XII-1930, Jaffuel 1306 (CONC); Cerros
de La Toma, 200 m, 30-XI-1934, Junge s.n. (CONC 5182);
Estero Membrillo, cerca de Agua de la Gloria, 150 m,
36º51’S, 72º54’W, 6-X-1980, R. Rodríguez 1476 (CONC);
Isla Quiriquina, 20 m, 27-XI-1924, Gunckel s.n. (CONC
116257); desembocadura del Bío-Bío, 5 m, 20-XII-1950,
Ricardi s.n. (CONC 10643); boca del Bío-Bío, lado Norte,
5 m, 9-XI-1935, Pfister s.n. (CONC 3557); la junta entre los
ríos Bío-Bío y Mininco, 110 m, 5-XII-1953, Sparre & Smith
80
80 (CONC); Caleta Burca, 36º28’S, 72º54’W, 7-XII-1980,
Ugarte 174 (CONC); Tomé Collen, 100 m, 24-XII-1934,
Junge s.n. (CONC 5253); Lirquen, Quebrada Honda, 100 m,
8-XII-1941, Behn s.n. (CONC 20872); Prov. Ñuble, camino
a Cobquecura, aproximadamente 10 km después del desvío
hacia El Boldo, 435 m, 36º07’S, 72º38’W, 6-XII-2001,
Mihoc et al. 3308 (CONC); Río Colmuyao interior, 35 m,
36º17’S, 72º47’W, XII-2001, Mihoc & Teneb 3897 (CONC).
Región de La Araucanía. Prov. Cautín, Temuco, 130 m, XI1946, Gunckel 43572 (CONC); Cunco (Huichahue), 200 m,
15-XII-1930, Aravena 271 (CONC); Toltén, 10 m, 21-III1942, Gunckel 12826 (CONC); Puente Loncotripai, a 15
km de Tirúa, 120 m, 38º24’S, 73º23’W, 21-II-2001, Parra &
Torres 782 (CONC). Región de Los Lagos. Prov. Chiloé, Isla
Grande de Chiloé, trayecto Piruquina a Alcaldeo de LlauLlao, cerca del Río Chilcán, 175 m, 42º23’S, 73º52’W, 14II-1984, Villagrán 5699 (CONC); Isla Quilán, entre Punta
Mala y Punta Abrigo, 20 m, 43º24’S, 74º15’W, 2-II-1986,
Villagrán & Leiva 7269 (CONC); Isla Guafo, trayecto desde
Caleta Samuel al Faro, 100 m, 43º36’S, 74º44’W, 25-II1986, Villagrán & Leiva 7545 (CONC); Prov. Llanquihue,
Petrohué, 210 m, 14-I-1946, Pfister s.n. (CONC 6012); Lago
Llanquihue, Ensenada, 60 m, 20-II-1944, Pfister 786, 4278
(CONC); ídem, La Poza, 100 m, 8-IV-1939, Looser 66209
(CONC); Prov. Osorno, Lliflén, Cerro Lliflén, 125 m, 23-II1958, C. Marticorena & Furet 62 (CONC); Lago Puyehue,
Isla Fresia, 250 m, I-II-1956, Levi Heins 2911 (CONC);
Prov. Palena, Hornopirén, 40 m, 41º58’S, 72º25’W, 2-I2002, Aedo 7351 (CONC). Región de Los Ríos. Prov.
Valdivia, Cerro Esmeralda, 120 m, 40º18’S, 73º27’W, II1958, Schlegel 1537 (CONC); Valdivia, Cno. Costanera, 10
m, 10-IV-1973, Montero 9005 (CONC); Playa de Mehuin,
20 m, 11-III-1964, Santos 154 (CONC); Mehuin, 20 m,
18-II-1976, Montero 10151 (CONC); Corral, 25 m, 28XI-1932, Gunckel 3324 (CONC); ídem, 25 m, 30-I-1933,
Gunckel s.n. (CONC 46204); Corral, Niebla, 25 m, 11-I1964, Gunckel 41862 (CONC); San Juan, 50 m, 8-II-1930,
Gunckel 1488 (CONC); Morro Gonzalo, 160 m, 23-I-1932,
Gunckel 2999 (CONC); ídem, 50 m, 6-XI-1936, Gunckel
19193 (CONC); Quebrada La Aguada, 25 m, 22-II-1932,
Gunckel 3196 (CONC); ídem, 50 m, 10-III-1936, Gunckel
28012 (CONC); San Carlos, 50 m, 10-X-1931, Gunckel
36408 (CONC). Dpto. La Unión, camino entre Hueicolla y
La Unión, km 2, 22-I-1971, Weldt-Rodríguez 946 (CONC);
Villarrica, Welturne, cataratas de Huilo-Huilo, 520 m, 27III-1980, Montero 11575 (CONC). Archipiélago Juan
Fernández, Isla Masatierra, 400 m, II-1965, Fuhrmann s.n.
(CONC 116172); ídem, 30 m, 33º38’S, 78º50’W, 10-XI1966, Angulo 105 (CONC); ídem, Quebrada de El Inglés,
400 m, 14-XII-1965, Muñoz & Sierra 7327 (CONC); ídem,
Punto San Carlos, Cumberland Bay, 26-XII-1965, Meyer
9684 (CONC); ídem, V. Colonial, 450 m, 25-II-1955, Sparre
121 (CONC); ídem, San Juan Bautista, 50 m, 33º37’S,
78º50’W, 13-I-1996, Swenson & Stuessy 399 (CONC);
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
ídem, between El Castillo and the electric generating plant
for the town, 7-II-1990, Stuessy & López 11449 (CONC);
ídem, La Vaquería, 20 m, 26-I-1984, Stuessy & Crawford
6436 (CONC); ídem, 10-100 m, 5-II-1990, López 11403
(CONC); ídem, in open areas along sea cliffs, 90 m, 15-I1984, Stuessy et al. 6201 (CONC).
4. Pseudognaphalium cymatoides (Kunze ex DC.) Anderb.,
Opera Bot. 104: 147. 1991. Gnaphalium cymatoides Kunze
ex DC., Prodr. 6: 225. 1838. TYPE: Chile. Andes, Concon,
4200 m, Dec., E. F. Poeppig s.n., Diar. 379 (lectotype,
here designated, P 00704542, = photo!; isolectotype, HAL
0111785, = photo!). Fig. 8.
Gnaphalium ulophyllum Hook. & Arn., Bot. Beechey Voy.
31. 1841, syn. nov. TYPE: Chile. Valparaíso, Playa Ancha
1832, T. C. Bridges 229 (lectotype, here designated, K
000500321, = photo!; isolectotype, E 00369125, = photo!)
Gnaphalium cymatoides Kunze ex DC. var. glabrum Walp.,
Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 19 (Suppl.
1): 276. 1843. TYPE: ‛Chile. Valparaiso’ (not seen).
Gnaphalium moelleri Phil., Anales Univ. Chile 90: 11. 1895.
Pseudognaphalium moelleri (Phil.) Anderb., Opera Bot.
104: 147. 1991, syn. nov. TYPE: Chile. Biobío, Renaico,
III-1885, R.A. Philippi s.n. (lectotype, here designated, SGO
64417!); isolectotypes K 000500403, =photo!, B (probably
destroyed), =photo 15109!
Gnaphalium canum Phil., Anales Univ. Chile 90: 11. 1895,
nom. illeg., non Wallich, 1831, syn. nov. TYPE: Chile.
Biobío, Renaico, III-1887, Philippi s.n. (lectotype, here
designated, SGO 64418!; isolectotypes, LP 001902!, LP
001903!, SGO 44965!).
Annual or biennial herbs, (7)10–40(50) cm tall,
multistemmed (rarely single stem), stems erect or
ascending, branched, weak, glandular-arachnoid. Leaves
9–35 × 1–2(6) mm, linear, margin revolute, conspicuously
undulate, apex acute, mucronate, base broad, scarcely
decurrent; concolorous, arachnoid-glandular on both
surfaces, trichomes of 2 types, short glandular biseriate
and few long eglandular eseptate, 3–4-celular trichomes.
Capitula in clusters arranged in lax corymbs or panicles.
Involucre campanulate, 2–3 × 2–3 mm. Phyllaries 3-seriate,
lamina whitish or yellowish, hyaline and shiny, apex obtuse;
outer phyllaries 2–3 × 0.8–1.5 mm, ovate; inner phyllaries
2.5–3 × 0.4–1 mm, oblong-obovate. Pistillate florets
40–95(130), corolla yellow, filiform, 1.5–3 mm. Bisexual
florets 4–8(11-12), corolla yellow, tubulose, 1.8–3 mm long.
Achenes 0.3–1 mm long, pilose with short oblong duplex
hairs. Pappus bristles 1.5–2.2 mm long.
PHENOLOGY. Flowering from (July, August) September to May.
DISTRIBUTION AND HABITAT. Bolivia (Hind 2011), Argentina
(Chubut, Neuquén, Río Negro) and North, Central and
South Chile (Arica y Parinacota, Coquimbo, Valparaíso,
Metropolitana, Libertador General Bernardo O´Higgins,
Maule, Biobío, Araucanía, Los Lagos), growing from sea
level to 3600 m.
RELATIONSHIPS. Pseudognaphalium cymatoides is closely
related to P. gayanum. Both species have undulate leaf
blade margins. Pseudognaphalium cymatoides can be
distinguished by its weakly branched stems and involucres
2–3 mm high (vs. robust unbranched stems and involucres
4–5 mm high in P. gayanum).
OBSERVATIONS
1. Candolle (1838: 225) mentioned two syntypes in the
protologue of Gnaphalium cymatoides: ‛In Chili circa
Concon (Poepp.!) in lapidosis maritimis secus torrentes
ad Valparaiso et S. Iago (Bert.!)’. We found these two
collections kept at P and MPU, Andes, 4200´ Decbr. et
ad Concon, (Diar. 379!!), Poeppig s.n., P 704542, and
Valparaiso: Prope St. Yago, ad follas et lepos, Bertero s.n in
3/1829, MPU 16284, respectively. We propose the material
‛Concon, Poeppig s.n.’, which presents the most complete
plant, as lectotype of Gnaphalium cymatoides.
2. We propose to place Gnaphalium ulophyllum (which
is characterized in the protologue by its linear leaves
arachnoid pubescent with undulate margins) into synonymy
of P. cymatoides because there is no diagnostic character
that differentiates the former from the latter.
3. According to the protologue, Gnaphalium ulophyllum
Hook. & Arn., was based on specimen ‘Valparaiso, Mr.
Bridges’. We located two sheets at E and K: E 00369125,
K 000500321. We selected the collection ‘K 000500321’,
which is better preserved.
4. Philippi (1895: 11) mentioned two syntypes in the
protologue of Gnaphalium moelleri, ‛In arvis inter Renaico
et Negrete in provincia Biobío’. We found these two
collections kept at SGO, [Negrete, III 85], SGO 44977
and [Renaico, Martio 1885], SGO 64417. We propose the
material SGO 64417, which presents the most complete
plant, as lectotype of Gnaphalium moelleri.
5. Gnaphalium moelleri Phil. is placed into synonymy of G.
cymatoides Kuntze ex DC. because there is no diagnostic
character that differentiates the former from the latter. Both
are glandular-arachnoid plants with linear leaves broad at
base with undulate margins and capitula in clusters arranged
in lax corymbs or panicles.
81
Gayana Bot. 71(1), 2014
FIGURE 8. Pseudognaphalium cymatoides. A, Habit; B, Leaf pubescence; C, D, Foliar trichomes; E, Capitulum; F, Phyllaries; G, Pistillate
floret; H, Bisexual floret. I, Distribution [A–H, Gay s.n., GH].
FIGURA 8. Pseudognaphalium cymatoides. A, Planta; B, Pubescencia de la hoja; C, D, Tricomas foliares; E, Capítulo; F, Filarios; G, Flor
pistilada; H, Flor perfecta. I, Distribución [A–H, Gay s.n., GH].
82
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
6. According to the protologue, Gnaphalium canum Phil.,
was based on specimen ‘Renaico, III-1887’. We located four
sheets at LP and SGO: LP 001902, 001903; SGO 64418,
44965. We selected as lectotype the collection ‘SGO 64418’
which is better preserved and has Cabrera’s annotation
‘Typus’.
7. Gnaphalium canum Phil. which was previously placed
into synonymy of G. moelleri Phil. (Cabrera 1971) is here
considered as synonym of Pseudognaphalium cymatoides.
ADDITIONAL MATERIAL EXAMINED. CHILE. Región de Arica
y Parinacota. Prov. Arica, Quebrada de Ipilla, 3000 m,
18º37’S, 69º28’W, 11-IX-1963, Schlegel 4895 (CONC);
Prov. Parinacota, cerros frente a Putre, 3600 m, 18º12’S,
69º34’W, 21-III-1987, Matthei & Rodríguez 269 (CONC).
Sin loc. determinada, 1838-1842, Herbarium of the U.S.
South Pacific Exploring Expedition under the command of
Capt. Wilkes, U.S.N (PH 01084668, GH 00282516); ídem,
Cuming 334 (SI). Región de Coquimbo. Prov. Choapa,
Illapel, 20-I-1942, Barros 2260 (SI); ídem, 1838-1842,
Philippi s.n. (NY); Illapel, Carretera Panamericana, 15
km N de Quilimarí, ca. Totoralillo, 25 m, 15-X-1971, C.
Marticorena et al. 1338 (CONC); Illapel, Caleta Oscuro, 5-50
m, 31º25’S, 71º35’W, 2-XI-1974, C. Marticorena et al. 276
(CONC); 2,1 km N of Puerto Oscuro on the Panamericana,
200 m, 31º24’S, 71º35’W, 18-IX-1994, Bliss & Lusk 575
(CONC); ídem, 20-XI-1994, Bliss 674 (CONC); Caleta
Manso, 9 km N de Huentelauquén, 20 m, 31º30’S, 71º34’W,
31-X-2000, Teillier & Márquez 4870 (CONC); Prov. Elqui,
Coquimbo, VII- VIII 1856, Harvey s.n. (GH 00282514); 16
km N of ‛Camping Morillos’ (84 km N of Los Socos) on rte.
5, 100 m, 11-II-1988, C. Marticorena et al. 9972 (CONC);
Carretera Panamericana entre Socos y Coquimbo, km 84,
Alto del Centinela, 100 m, 30º01’S, 70º23’W, 11-II-1988,
C. Marticorena et al. 9972 (CONC); Andacollo, 11-X-1958,
Ricardi & C. Marticorena 4943 (CONC); ídem, Ricardi &
C. Marticorena 4943 bis (CONC); Prov. Limarí, Ovalle,
Tulahuén, 15-I-1942, Barros 2255 (SI); Ovalle, Talinay,
700 m, 30º50’S, 71º37’W, 4-III-1950, Jiles 1686 (CONC);
Ovalle, Río Limarí, 180 m, 30º37’S, 71º12’W, 16-XII-1948,
Jiles 1112 (CONC). Región de Valparaíso. Prov. Los Andes,
Portillo, Laguna del Inca, 3200 m, 32º50’S, 70º09’W, 11-III1994, Teillier & Niemeyer 3322 (CONC); Prov. Quillota, La
Cruz, Fundo Sta. Ana, 100 m, 22-IV-1945, Behn s.n. (CONC
20871); Prov. Petorca, Zapallar, 35 m, 15-XI-1928, Behn s.n.
(CONC 20870); Zapallar, Cerro Francés, 600 m, 23-II-1953,
Levi Heins 150 (CONC); ídem, 20-II-1953, Levi Heins 174
(CONC); Prov. San Antonio, Palos Quemados, 22-I-1933,
Looser 66226 (MO); Prov. San Felipe de Aconcagua, Jahuel,
El Zaino, 1100 m, 11-XI-1955, Barrientos 1699 (CONC);
Prov. Valparaíso,Viña del Mar, 21-II-1939, Burkart 9383
(SI); Quintero, 40 m, XII-1950, Gunckel 21612 (CONC);
Quintero, Los Juanes, 20 m, IX-1953, Navas s.n. (CONC
116177 y 116178); ídem, 20-I-1954, Navas s.n. (CONC
116326); Quintero, Dunas de Ritoque, 10 m, 32º46’S,
71º32’W, II-1963, Gunckel 40170 (CONC); Prov. MargaMarga, Valle de Marga-Marga, IX-1932, Jaffuel & Pirion
3026; ídem, 1930-32, Jaffuel & Pirion 3226 (CONC);
Marga-Marga, 150 m, I-1914, Jaffuel s.n. (COvillagránNC
89741); S/Prov., Laguna Peñuelas, 350 m, 27-II-1952, s/leg.
(CONC 13243); Tranque Orosco, 290 m, V-1966, Zöllner
1068 (CONC). Región Metropolitana de Santiago. Prov.
Santiago, Santiago, s .f., M. Cl. Gay s.n. (GH); Quebrada de
La Plata, 740 m, 33º29’S, 70º54’W, 23-XI-1960, Schlegel
3199 (CONC); ídem, laderas de los cerros Morro El Fraile
y Buitreras, 540-630 m, 33º29’S, 70º50’W, 4/18-II-2001,
Mihoc 4 (CONC); Santuario de la Naturaleza Yerba Loca,
vega en ladera al SW del estero Yerba Loca, 1880 m, 33º20’S,
70º19’W, 31-I-1999, Arroyo & Humaña 99887 (CONC); El
Arrayán, 14-I-1954, Arriagada s.n. (CONC 116277); Cerro
San Cristóbal, 800 m, 14-V-1954, Navas 632 (CONC).
Región del Libertador General Bernardo O´Higgins.
Prov. Cachapoal, Rancagua, 2-I-1938, Barros 2307 (SI);
Prov. Colchagua, Río Claro, La Rufina, 315 m, I-II-1959,
Barrientos 2025 (CONC); S/Prov., Las Peñas, 755 m, 25I-1958, Barrientos 1621 (CONC). Región del Maule. Prov.
Linares, Camino Parral-Cauquenes, km 10, Parcela 2, El
Cairo, 150 m, 36º06’S, 71º54’W, 17-III-1988, Rodríguez &
Baeza 2371 (CONC); Prov. Talca, Los Bellotos del Melado,
700-800 m, 35º50’S, 71º06’W, 1-III-2005, Mihoc 60
(CONC). Región del Biobío. Prov. Biobío, Negrete, a orillas
del Bío-Bío, 75 m, 19-II-1951, Pfister s.n. (CONC 10503);
Prov. Concepción, Hualpén, 20 m, 9-III-1940, Junge s.n.
(CONC 6771); Prov. Ñuble, Quirihue, 200 m, 12-IV-1952,
Zemelman s.n. (CONC 13277); camino a Recinto, 950 m,
26-IV-1954, Ledezma 8 (CONC). Región de La Araucanía.
Prov. Cautín, Temuco, ca. Río Cautín, 110 m, 14-II-1941,
Gunckel 11011 (CONC); Temuco, Truf Truf, 125 m, 20-XII1960, Montero 6364 (CONC); Almagro, 50 m, 21-II-1943,
Gunckel s.n. (CONC 116238); Cajón, 130 m, 26-III-1946,
Cañulaf s.n. (CONC 116241); Toltén, lado N desembocadura
Río Toltén, 10 m, 21-III-1942, Gunckel 12827 (CONC);
Volcán Villa-Rica, 1300 m, I-1926, Hollermayer s.n. (CONC
116282); Prov. Malleco, Angol, peddly gravelly shore of Río
Malleco, 100-150 m, 27-28-II-1925, Pennell 12845 (PH).
Región de Los Lagos. Prov. Osorno, Ribera N del Lago
Rupanco, Fundo Pichilafquen, 120 m, 13-II-1955, Pfister
s.n. (CONC 18562); Pichilafquén-Lago Rupanco, 120 m,
15-II-1956, Pfister s.n. (CONC 19025).
5. Pseudognaphalium gayanum (J. Rémy) Anderb., Opera
Bot. 104: 147. 1991. Gnaphalium gayanum J. Rémy in
Gay, Fl. Chilena 4(3): 225. 1849. TYPE: Chile. Colchagua,
Cordillera de San Fernando, Talcaregue, II-1831, C. Gay
s.n. (holotype, P 00704552, = photo!). Fig. 9.
83
Gayana Bot. 71(1), 2014
FIGURE 9. Pseudognaphalium gayanum. A, Habit; B, Leaf pubescence; C, D, Foliar trichomes; E, Capitulum; F, Phyllaries; G, Pistillate
floret; H, Bisexual floret. I, Distribution [A–H, Werdermann 470, NY].
FIGURA 9. Pseudognaphalium gayanum. A, Planta; B, Pubescencia de la hoja; C, D, Tricomas foliares; E, Capítulo; F, Filarios; G, Flor
pistilada; H, Flor perfecta; I, Distribución [A–H, Werdermann 470, NY].
84
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
Gnaphalium heterotrichum Phil., Linnaea 29: 4. 1857-58.
Pseudognaphalium heterotrichum (Phil.) Anderb., Opera
Bot. 104: 147. 1991, syn. nov. TYPE: Chile. Coquimbo,
La Serena, X-1837, C. Gay 681 (holotype, SGO 71285!;
isotype, B (probably destroyed), = photo F15100 SI!),
related to P. cymatoides. Both species have conspicuously
undulate leaf blade margins. Pseudognaphalium gayanum
can be distinguished by its robust unbranched stems and
involucres 4–5 mm high (vs. weakly branched stems and
involucres 3–4 mm high in P. cymatoides).
Gnaphalium leucocephalum Phil., Linnaea 33: 166.
1864, nom. illeg., non A. Gray, 1853. Gnaphalium
pseudohelichrysum Reiche, Anales Univ. Chile 112: 112.
1903, nov. nom., syn. nov. TYPE: Chile. Colchagua, 2-3000
m s.m., XII-1860, Landbeck s.n. (holotype, SGO 64445!;
isotype, B probably destroyed).
OBSERVATIONS
1. Muñoz Pizarro (1960: 142) cited as holotype of
Gnaphalium heterotrichum the collection ‛cerca de La
Serena, Gay. Herb, Chil. Nº 681, 64406’. However at SGO
was located the collection ‛Prov. Coquimbo, Serena, 8bre.
1837, [Gay] 681, 71285’. Probably ‛64406’ instead of
‛71285’ was a transcription error.
Gnaphalium heterophyllum Phil., Anales Univ. Chile 90:
14. 1895, syn. nov. TYPE: Chile. Coquimbo, year 1888,
W. Geisze s.n. (lectotype, here designated, SGO 64403!;
isolectotypes, LP 001921!, SGO 64402!, 64404!, 72256!)
Gnaphalium robustum Phil., Anales Univ. Chile 90: 16.
1895. Pseudognaphalium robustum (Phil) Anderb., Opera
Bot. 104: 148. 1991, syn. nov. TYPE: Chile. Santiago, Rio
Colorado, I- 1888, Philippi s.n. (holotype, SGO 64424!).
Perennial herbs, 30–110 cm tall, multistemmed, stems erect
or ascending, unbranched, robust, arachnoid-glandular
to whitish-woolly. Leaves (30)40–60(90) × 2–4(10)
mm, linear, margins conspicuously undulate, apex acute,
mucronate, base short-decurrent 5-11 mm long, sometimes
broad; concolorous, glandular-arachnoid, trichomes of 2
types on both surfaces, glandular biseriate and few long
eglandular eseptate, 5-celular trichomes. Capitula in clusters
arranged in corymbs or panicles (seen in large plants).
Involucre broadly campanulate 4–5 × 4–5 mm. Phyllaries
4-seriate, lamina light-brown, yellowish, hyaline and shiny
or opaque white, apex obtuse to rounded; outer phyllaries
ca. 5 × 2 mm, ovate; inner phyllaries ca. 3.5 × 1 mm, oblongobovate. Pistillate florets 84–181, corolla yellow, filiform,
2–2.6(3) mm long. Bisexual florets (7)13–17(31), corolla
yellow, tubular, 2–3.6 mm long. Achenes 0.1–0.3 mm long,
pilose with short oblong duplex hairs. Pappus bristles 2.2–
3.4 mm long.
PHENOLOGY. Flowering from (May, July) September to April.
VERNACULAR NAME.‛Té de burro’.
DISTRIBUTION AND HABITAT. Endemic to Chile from northern,
central and southern Regions (Arica y Parinacota, Atacama,
Coquimbo, Valparaíso, Metropolitana, Libertador General
Bernardo O´Higgins, Maule, Biobío, Los Lagos, Aysén
del General Carlos Ibañez del Campo, Magallanes y de la
Antártica), growing from sea level to 3500 m.
RELATIONSHIPS. Pseudognaphalium gayanum is closely
2. Gnaphalium heterotrichum Phil. was described on
the basis of its white tomentose stems. However, this is
a variable feature throughout the range of Gnaphalium
gayanum J. Rémy.
3. Gnaphalium leucocephalum Phil. (= G. pseudohelichrysum
Reiche, nov. nom.) was described on the basis of its
white-yellowish phyllaries and white tomentose leaves.
However, these are variable features throughout the range
of Gnaphalium gayanum J. Rémy.
4. There is a photo-Field Museum Nr.15107 ex B, with a
specimen collected by Philippi in Cordillera de Linares.
Since the protologue of Gnaphalium leucocephalum
mentions ‘Landbeck, Colchagua’ this specimen is excluded
from the type collection of G. leucocephalum.
5. According to the protologue, Gnaphalium heterophyllum
Phil. was based on the specimen ‘Frequens in provincia
Coquimbo’. We located four sheets at SGO: ‘Coquimbo’,
‘Coquimbo, 10. 1878’, ‘Coquimbo, sept. 1888’ and
‘Coquimbo, W. Geisze, 1888’. We selected as lectotype
the collection ‘Coquimbo, W. Geisze, 1888’ which is better
preserved.
6. Gnaphalium heterophyllum Phil. which was previously
placed into synonymy of G. heterotrichum Phil. (Freire
& Iharlegui 2008) is here considered as synonym of
Pseudognaphalium gayanum.
7. After examining the type material, we propose to place
Gnaphalium robustum Phil. into synonymy of G. gayanum
J. Rémy, because there is no character that differentiates the
former from the latter. Both are multistemmed, arachnoidglandular to whitish-woolly plants with unbranched woody
stems, and linear leaves, undulate on margins.
ADDITIONAL MATERIAL EXAMINED. CHILE. Región de Arica y
Parinacota. Prov. Arica, camino de Arica a Portezuelo de
Chapiquiña km 96, 3450 m, 9-II-1964, C. Marticorena et al.
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Gayana Bot. 71(1), 2014
46 (CONC); Prov. Parinacota, Camino a Putre, 3200 m,
18º12’S, 69º34’W, 11-V-2000, Belmonte 20154 (CONC);
camino Zapahuira a Putre, km 13, 3300 m, 5-V-1972, Ricardi
et al. 155 (CONC). S/Región, Río Blanco, 2000 m, 27-XII1964, Zöllner 1076 (LP). S/loc., 13-VI-1948, Bultmann
23661 (CONC); s/loc., s/fecha, Hno. C. Joseph 3657
(CONC). Región de Atacama. Prov. Copiapó, Totoral, ca.
100 m, II-1924, E. Werdermann 470 (LP, MO, NY, SI); Prov.
Huasco, 1 km S of Huasco, 20-X-1955, Böcher et al. 543
(LP); Quebrada del Pretil, 14-XI-1956, Ricardi & C.
Marticorena 3956 (CONC); Carrizal Bajo, Mina Oriente,
320 m, 28º08’S, 71º07’W, 14-XI-1985, Schlegel 8034
(CONC). Región de Coquimbo. Prov. Choapa, Illapel, s. f.,
Philippi s.n. (NY); ídem, 310 m, 11-IX-1924, Barros 3404
(CONC); ídem, 310 m, IX-1926, Barros 1467 (CONC);
Illapel, Cuesta del Espino, 3-X-1941, Barros 2318 (LP);
Caren, near Quebrada Luncuman, ca. 40 km from Illapel,
tributary Illapel river., ca. 1000 m, 14-XI-1938, Worth &
Morrison 16494 (MO, SI); Caleta Manso, 9 km N
Huentelauquén, 20 m, 31º30’S, 71º34’W, 31-X-2000,
Teillier & Márquez 4848 (CONC); Valle Ojotas, cordillera
Quelen, cerca de Cuncumen, 2500 m, I-1984, Zöllner 11790
(SI); Prov. Elqui, Cuesta Buenos Aires nördl La Serena, 29º
34´S, 71º 15´W, 500 m, 11-XII, 1987, Rosas 1385 (M);
Cuesta Buenos Aires, 500 m, 29º34’S, 71º14’W, 31-X-1991,
Rodríguez 2789 (CONC); Cuesta de Buenos Aires, cerca del
Portezuelo, 550 m, 20-X-1971, C. Marticorena et al. 1612
(CONC); entre Coquimbo y Buyarán, 100 m, 8-XI-1923,
Werdermann 107 (SI); Coquimbo, de La Serena a Ovalle, 41
km sur de La Serena, cuesta las Cardas, 30º 15´S, 71º 17´W,
450 m, 14-XII-1987, Rosas 1507 (M); Coquimbo 30ºS, 2030 m, 13-10-1898, Von Bayern 388 (M); Coquimbo, Playa,
15 m, 26-IX-1934, Montero 1881 (CONC); Punta de
Coquimbo, 20-IX-1957, Ricardi & C. Marticorena 4488
(CONC); ídem, 20-IX-1957, Cabrera 12697 (LP); vicinity
of Coquimbo, IX-1931, Jaffuel 2681 (GH); Quebrada al
Norte del Punto Juan Soldado, 29º 39´S, 71º 18´W, 11-XII1987, Rosas 1392 (NY); Quebrada El Pangue al N de
Cochiguaz y El Colorado 30º 07´S, 70º 23 W, 2000 m, 13XII-1897, Rosas 1465 (M); Andacollo, 1000 m, 11-X-1958,
Ricardi & C. Marticorena 4972 (CONC, LP); Paihuano, III1946, Roesner s.n. (CONC 6494); Paihuano, cerca del río,
950 m, 25-IX-1948, Behn s.n. (CONC 8414 y 20882);
Chanchoquí-Paihuano, 900 m, 20-IX-1948, Pfister s.n.
(CONC 8363); Along the Panamerican Highway, 20 km S of
Las Herraduras, 400 m, 30º05’S, 71º25’W, 27-I-1991, De
Vore 1545 (CONC); Camino de Playa Temblador a Cruz
Grande, 198 m, 29º28’S, 71º17’W, 17-IX-1989, Niemeyer et
al. 89122 (CONC); Cochiguaz, 26-V-1990, Von Bohlen 742
(CONC); La Serena, IX-1928, Barros 24725 (CONC); Punta
Teatinos, 10 km N de La Serena, 15-X-1940, Looser 4394
(LP); puente Las Trancas, en el camino de Vicuña a Hurtado,
16-XII-1940, Wagenknecht 4396 (LP); Rivadavia, cauce del
Río Turbio, 17-I-1936, Cabrera 3506 (LP); Prov. Limarí,
86
Altos de Talinay, 700 m, 18-XI-1978, Zöllner 10442 (2
ejemplares, MO, CONC); Talinay, 700 m, 30º50’S, 71º37’W,
8-II-1948, Jiles 542-a (CONC); Cerro Talinay, 550 m, 8-XII1953, Kausel 3806 (LP); Cordillera de Ovalle, Río Tascadero,
El Polvo, 31º08´S, 70º35´W, 2400 m, Jiles 6437 (M); Ovalle,
Fray Jorge, X-1951, Jiles s.n. (CONC 89696); Cordillera de
Ovalle, Quebrada Larga, 3000 m, 30º44’S, 70º23’W, 3000
m, 17-II-1958, Jiles 3429 (CONC); Hacienda Tamaya, 250
m, 30º34’S, 71º24’W, 30-X-1956, Jiles 3047 (CONC);
Estancia El Parral, 600 m, 21-IX-1950, Jiles 1848 (CONC,
LP); Camino de La Serena a Ovalle por Las Cardas, km 61,
pasando la Cuesta Las Cardas, Quebrada del Ingenio, 3,5 km
S de Estación Higueritas, 380 m, 30º26’S, 71º10’W, 11-II1988, C. Marticorena et al. 9956 (CONC); Quebrada del
Teniente, 30 m, 9-II-1963, Ricardi et al. 790 (CONC);
Carretera Panamericana, 11 km S de la Quebrada Teniente,
9-II-1963, Ricardi et al. 796 (CONC); Along the Panamerican
Highway, 1 km N of Mantos de Hornillo, 250 m, 17-I-1989,
Lammers et al. 6381 (CONC); Parque Nacional Fray Jorge,
500 m, 30º40’S, 71º40’W, 5-XI-1974, C. Marticorena et al.
468 (CONC); ídem, 450 m, 15-IX-1947, Ibáñez et al. s.n.
(CONC 144832); ídem, X-1951, Jiles s.n. (CONC 89696);
ídem, 17-VII-1967, Zöllner 1772 (LP); S/Loc., 9-IX-1926,
Barros 40 (LP); S/loc., 3-X-1953, Cabrera 11415 (LP).
Región de Valparaíso. Prov. Los Andes, Uspallata-Pass der
chilenischen Hochcordillere (33ºS Breite): Juncal; zwischen
Geröll, 2200 m, 20-I-1903, Buchtien s.n. (SI 022607);
Juncal, 2200 m, II-1903, Buchtien 3 (MO); Cajón del Río
Colorado, 3000 m, 32º34’S, 70º20’W, 20-II-2002, Teillier
5111 (CONC); Saladillo, Laguna Turquesa, Río Estero
Castro, 3560 m, 33º06’S, 70º16’W, 17-IV-2007, Rosas 4799
(CONC); Portillo, 2800 m, I-1970, Crisci 490 (LP); Prov.
Marga Marga, Limache, 90 m, I-1948, Levi Heins 445
(CONC); Estero de Maitenes, Maitenes, Limache, 27-IX1928, Garaventa 320 (CONC); Cordillera de la Costa, cerca
de Limache, en un cerro, Punta Imán, 1800 m, XI 1964,
Zöllner 978 (LP); Cerro La Campana, 1200 m, 17-XII-1973,
Zöllner 8088 (CONC, MO); Cerro de La Campana mirando
a Olmué, 500-1000 m, 22-XI-1936, Looser 3765 (CONC);
Cerro Campana, cerca de la mina ‛La Pronosticada’, 1300
m,19-I-1936, Garaventa 3117 (CONC); cerca de Quilpué,
XI-1967, Zöllner 2200 (LP); Prov. Petorca, Rt. 5 between
tunnel and La Ligua, 110 m, 32º33’25’’S, 71º15’54’’W,
3-XI-2006, Tepe et al. 1688 (CONC); La Ligua, Quebrada
Las Pataguas, 160 m, XII-1958, Escudero s.n. (CONC
75387); Las Palmas de Pedegua, 600 m, X-1945, Moreno
s.n. (CONC 116360); Aguas claras cerca de Zapallar, 25 m,
11-II-1953, Levi Heins 175 (CONC); Prov. Quillota, Cerro
de la Campana Chica, 29-XII-1937, Barros 195, 2321 (LP);
Cerro Roble above Calco ca. 2 km by trail, 1210 m, 2-XII1951, Hutchinson 67 (GH); Embalse de Los Aromos, 100 m,
32º56’S, 71º20’W, 10-XI-2001, Aedo 6806 (CONC); Cuesta
El Melón, 560 m, 32º40’S, 71º15’W, 28-VII-1982, Bayer 42
(CONC); ídem, 13-IX-1957, Ricardi & C. Marticorena 4223
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
(CONC); ídem, XI-1969, G. Martínez 49072 (CONC); cerca
de La Calera, Cordillera de la Costa, 600 m, 13-XI-1966,
Zöllner 1138 (LP); Prov. San Antonio, Rocas de Santo
Domingo, 25 m, 24-VII-1955, Gunckel s.n. (CONC 116263);
Prov. San Felipe de Aconcagua, Carretera Panamericana,
Bajada de El Melón, 13-IX-1957, Cabrera 12528 (LP, M); El
Melón, 560 m, II-1953, Collantes s.n. (CONC 75383);
Portillo, 2870 m, 6-III-1954, Ricardi 2853 (CONC); Santa
Filomena, al interior de Jahuel, 900 m, 32º41’S, 70º37’W,
13-XI-1987, Matthei & Quezada 463 (CONC); Prov.
Valparaíso, Bajada Agua Santa, 9-I-1950, Boelcke 3885 (LP,
SI); Las Rocas, 26-XII-1949, Boelcke 3846 (LP, SI);
Casablanca, 18-XI-1954, Navas 1389 (CONC); Tunquén,
estero Casablanca, 10-50 m, 33º16’S, 71º39’’W, 1-XI-2001,
Teillier 5851 (CONC); ca. Placilla de Peñuelas, 360 m, 16XI-1954, Sparre 11016 (CONC); between Maitencillo and
Puchuncaví, 26-X-1985, Zöllner 12518 (MO); MargaMarga, 33º06’S, 71º25’W, IX-1931, Jaffuel 3267 (CONC);
Viña del Mar, 20 m, XI-1953, Gunckel 32025 (CONC); Viña
del Mar, El Salto, 110 m, 23-XI-1958, Saa s.n. (CONC
75390); Viña del Mar, Fundo Siete Hermanas, 270 m, 4-XI1936, Behn 24413 (CONC); Recreo Alto, 25 m, 24-XI-1958,
Parra 475 (CONC); Granizo, 300 m, 25-X-1959, Weisser
Sievers s.n. (CONC 75846); Pucalán, 31-XII-1950, Collantes
s.n. (CONC 75384); Cerro Curauma, 33º10’5,3’’S,
71º41’4,86’’W, 20-I-2006, Fernández 1494 (CONC); Valle
del agua potable, Las Lúcumas, 60 m, 14-XI-1954, Schlegel
433 (CONC); S/Prov., Punta Cristales, Cerro Roble, 1800 m,
29-XI-1965, Zöllner 1069 (CONC). Región Metropolitana
de Santiago. Prov. Chacabuco, Panamericana Norte km 22,
Straβenrand mit nassem Graben., 3 XII 1966, Zollitsch 22
(M); Prov. Cordillera, inmediaciones Refugio Lo Valdés,
Valle de Maipo, 2000 m, 20-XI-1940, Garaventa 1293
(CONC); Cajón del Maipo, Refugio Cruz de Piedra, 34º06’S,
70º03’W, 2400 m, 20-I-2000, Teillier 4624 (CONC); Prov.
Maipo, Cajón de Morales entre las Panimávidas y la laguna,
2200 m, 33º47’S, 70º04W, 25-I-2002, Teillier & Márquez
5313 (CONC); ídem, 2150 m, Teillier & Márquez 5314
(CONC); Laguna de Aculeo, 350 m, 18-V-1971, C. Martínez
s.n. (CONC 74364 y 75845); ídem, 11-X-1974, Montero
9437 (CONC); Laguna Negra, ribera oriental, 2720 m,
33º00’S, 69º02’W, 15-I-1990, Teillier et al. 2027 (CONC);
Valle del Yeso, Estero Salinillas, 2900 m, 33º40’S, 69º57’W,
IV-2001, Teillier 6489 (CONC); Prov. Santiago, entre
Rungui y La Capilla (Calen), 700-1000 m, 6-XI-1937,
Looser 66253 (MO); Comuna de la Reina, Quebrada de
Ramón, 800-1800 m, 33º26’S, 70º30’W, 21-I-2001, Tomé
128, 129 (CONC); Valle Macul, 1850 m, XII-1932, Grandjot
995 (CONC); Quebrada de La Plata, 740 m, 33º29’S,
70º54’W, 15-XII-1960, Schlegel 3329 (CONC); S/Prov.,
Cuesta de La Dormida, 1200 m, IX-1956, Gunckel 30004
(CONC); ídem, 13-I-1972, Mahu 8657 (LP). Región del
Libertador General Bernardo O´Higgins. Prov. Cachapoal,
Hacienda de Cauquenes, Cajón del Cypres, 1875, Dessauer
s.n. (M 0155925); Reserva Nac. Los Cipreses, 1062 m,
34º15’S, 70º25’W, 2-I-2006, A. Marticorena & Jiménez s.n.
(CONC 173679); Pangal, 1000 m, 25-XI-1978, Zöllner
10151 (CONC); Termas de Cauquenes, camino a Coya, 800
m, 4-XI-1952, Pfister s.n. (CONC 13160); Prov. Colchagua,
Laguna Cahiul, 50 m, 12-II-1954, Arriaga s.n. (CONC
75391). Región del Maule. Prov. Curicó, La Montaña, Teno,
27-I-1945, Barros 3874 (CONC); Valle Vergara, 920 km del
límite entre Chile- Argentina, II-1967, Calderón s.n. (CONC
34579). Región del Biobío. Prov Ñuble, Puente El Roble, 64
m, 36º45’S, 72º25’W, 14-XI-1972, Igaymán & M. Muñoz
s.n. (CONC 89695). Región de Los Lagos. Prov. Llanquihue,
Petrohué, 210 m, 18-I-1979, Zöllner 10198 (CONC). Región
de Aysén del General Carlos Ibáñez del Campo. Prov.
Coyhaique, Lago Caro, 285 m, 45º48’S, 72º34’W, 23-II2007, García 4083 (CONC). Región de Magallanes y de la
Antártica Chilena. Prov. Última Esperanza, Puerto Prat,
Seno Última Esperanza, 26-I-1904, Hicken 122 (SI).
6. Pseudognaphalium lacteum (Meyen & Walp.) Anderb.,
Opera Bot. 104: 147. 1991. Gnaphalium lacteum Meyen &
Walp., Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 19
(Suppl. 1): 276. 1843. TYPE: Perú. Meyen s.n. (holotype,
B (probably destroyed), = photo F15105!; lectotype, here
designated, GH 00008360, = photo! fragment ex B). Fig. 10.
Gnaphalium argyrolepis Phil., Anales Mus. Nac., Santiago
de Chile 8: 46. 1891. TYPE: Chile. Iquique. Usmagama, 15III-1885, C. Rahmer s.n. (lectotype, here designated, SGO
64432!).
Dwarf perennial herbs, 2–8 cm tall, multistemmed, stems
prostrate, branched, whitish woolly. Leaves approximate,
ca. 15 × 3 mm, obovate, margin flat, apex rounded, base
atenuate; concolorous, woolly on both surfaces, trichomes
of 2 types, long eglandular eseptate, 4-celular, and few
biseriate glandular trichomes hidden under the wool.
Capitula few, in clusters arranged in corymbs or panicles.
Involucre broadly campanulate, ca. 5 × 5 mm. Phyllaries
4-seriate, lamina usually opaque milky-white, rounded;
outer phyllaries ca. 4 × 2–2.7 mm, broadly elliptic; inner
phyllaries 4–4.6 × 0.7–1 mm, oblong-lanceolate. Pistillate
florets 42–59, corolla yellow, filiform, ca. 3.8 mm long.
Bisexual florets 8-15, corolla yellow, tubular, ca. 3.7 mm
long. Achenes 0.6–1 mm long, glabrous, epidermis smooth.
Pappus bristles ca. 3.2. mm long.
PHENOLOGY. Flowering from September to May.
DISTRIBUTION AND HABITAT. Perú (Dillon & Sagástegui 1991),
Bolivia (Hind 2011), NW Argentina (Catamarca, Jujuy,
Salta) and northern Chile (Arica y Parinacota, Tarapacá,
Coquimbo, Biobío), between 3000-4800 m.
87
Gayana Bot. 71(1), 2014
FIGURE 10. Pseudognaphalium lacteum. A, Habit; B, Cluster of capitula and leaves; C, D, Foliar trichomes; E, Capitulum; F, Phyllaries; G,
Distribution [A, B, E, from Cabrera 1978; C, D, Werdermann 1092, LP; F, Antoniz s.n., CONC].
FIGURA 10. Pseudognaphalium lacteum. A, Planta; B, Detalle de la inflorescencia; C, D, Tricomas foliares; E, Capítulo; F, Filarios; G,
Distribución [A, B, E, tomado de Cabrera 1978; C, D, Werdermann 1092, LP; F, Antoniz s.n., CONC].
88
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
RELATIONSHIPS. Pseudognaphalium lacteum appears to
occupy an isolated position among Chilean species taking in
consideration its dwarf habit with prostrate branched stems
and usually opaque milky-white phyllaries.
OBSERVATIONS
1. Because the collection mentioned in the protologue of
Gnaphalium lacteum, ‛Peruvia: in planitie circa Tacoram, alt
14-17000 ped.’ and deposited at B was probably destroyed,
the isotype at GH (fragment), which is sufficiently
diagnostic, is designated as lectotype.
2. Two specimens from Tarapacá are mentioned in the
protologue of Gnaphalium argyrolepis: ‛Habitat in
provincia Tarapacá ad Usmagama, Vilon, etc.’. We found
at SGO three sheets, two from Vilon: [Vilon, pr. Atacama,
17-II-1885, Philippi s.n.] SGO 44934, 71282, and the other
from Usmagama, [Usmagama, 15-III-1885, C. Rahmer
s.n.] SGO 64432. We propose the material SGO 64432,
which presents the most complete plant, as lectotype of
Gnaphalium argyrolepis.
ADDITIONAL MATERIAL EXAMINED. CHILE. Región de Arica
y Parinacota. Prov. Arica, Cordillera de Arica, Cuesta de
Chapiquiña, cerca del portezuelo, 4000 m, 29-III-1962,
Ricardi et al. 325 (CONC); camino de Arica al Portezuelo
de Chapiquiña, km 92, 3250 m, 8-II-1964, C. Marticorena
et al. 14 (CONC); FF.CC. de Arica-La Paz, Puquios, km
106-108, 3600-3700 m, 27-IX-1955, Ricardi & Silva 3544
(BAB, CONC). Prov. Parinacota, cerca de Parinacota,
4400 m, 18º13’S, 69º14’W, 8-III-1984, Arroyo 84-691
(CONC); Putre, 3500 m, 12-II-1964, C. Marticorena et al.
142 (CONC); camino Zapahuira-Putre, Quebrada Chilcana,
3400 m, 18º14’S, 69º33’W, 15-III-1985, Landero et al. 48
(CONC); Lago Chungará, arriba guardería CONAF, 4600
m, 18º14’S, 69º10’W, 18-V-1989, Niemeyer et al. 89106
(CONC); Guaneguane, bofedal. 4450 m, 18º10’S, 69º15’W,
Niemeyer et al. 89101 (CONC); Lagunas de Cotacotani,
4500 m, 13-II-1964, C. Marticorena et al. 244 (CONC);
camino de Chucuyo a las Lagunas de Cotacotani, km 5, 4400
m, 13-II-1964, C. Marticorena et al. 227 (CONC). Región
de Tarapacá. Prov. El Tamarugal, Cerro Japu, 4200 m, III1926, Werdermann 1092 (CONC, NY, LP, SI); Tranque de
Caritaya, 3600 m, 30-IX-1958, Ricardi & C. Marticorena
4807 (CONC); camino de Huara a Cancosa, km 91, 3850
m, 17-II-1964, C. Marticorena et al. 316 (CONC). Vilon,
pr. Atacama, 17-II-1885, Philippi s.n. (SGO 44934, 71282).
Región de Coquimbo. Prov. Limarí, Carretera Panamericana,
19 km al S de Socos, 300 m, 9-II-1963, Ricardi et al. 788
(CONC); Cordillera de Combarbalá, Potrero Grande, 2800
m, 31º10’S, 70º50’W, 11-I-1973, Jiles 6083 (CONC).
Región del Biobío. Prov. Concepción, San Pedro, X-1952,
Antoniz s.n. (CONC 13265).
7. Pseudognaphalium landbeckii (Phil.) Anderb., Opera
Bot. 104: 147. 1991. Gnaphalium landbeckii Phil., Linnaea
33: 165. 1864. TYPE: CHILE. Colchagua, Llico, XII-1861,
Lanbeck s.n. (lectotype, here designated, SGO 64383!;
isolectotypes, LP 116929!, SGO 44952!). Fig. 11.
Perennial herbs, 15–30 cm tall, stem solitary (2 or 3), erect,
unbranched, whitish woolly. Stems leaves 35-40 × 5-7 mm,
broadly oblong, margin flat, apex obtuse, base decurrent 6-8
mm long; basal leaves 50-70 × 9-10 mm, linear-obovate;
concolorous, woolly, trichomes of 2 types, long eglandular
eseptate trichomes, 3–4-celular, and few biseriate glandular
trichomes hidden under the wool. Capitula numerous in
clusters arranged in dense corymbs. Involucre campanulate,
4–5 × 5 mm. Phyllaries ca. 42, 4-seriate, lamina brown,
hyaline and shiny; outer phyllaries 3–4 × 1.5–2.7 mm,
obovate, apex subobtuse, lanuginose; inner phyllaries 3.1–
4.6 × 0.7–1.3 mm, linear, apex acute, glabrous. Pistillate
ca. 180, corolla yellow, filiform, ca. 2.5 mm long. Bisexual
florets ca. 14, corolla yellow, tubular, ca. 2.7 mm long.
Achenes, 0.6–0.7 mm long, glabrous, epidermis smooth or
papillose. Pappus bristles 3–3.2 mm long.
PHENOLOGY. Flowering from October to December
(February).
DISTRIBUTION AND HABITAT. Endemic to Chile (Valparaíso,
Metropolitana, Maule), between 50-700 m.
RELATIONSHIPS. Pseudognaphalium landbeckii seems to
be distinguished by its usually solitary stems, numerous
capitula in clusters arranged in dense corymbs, and stem
leaves with broadly oblong leaf blades.
OBSERVATION
There is a photo-Field Museum Nr.15106, with a specimen
collected in Chillán. Since the protologue of Gnaphalium
landbeckii mentions ‘Llico’ this specimen is excluded from
the type collection of G. landbeckii.
ADDITIONAL MATERIAL EXAMINED. CHILE. Región de
Valparaíso. Prov. Valparaíso, Concón, 7-X-1953, Cabrera
11462 (LP). Región Metropolitana de Santiago. Prov.
Santiago, Clarillo, X-1958, Gunckel 46477 (CONC).
Región del Maule. Prov. Curicó, Llico, II-1950, Ojeda
20.949 (CONC); Prov. Talca, Constitución, Playa grande,
1-XI-1958, Barnier 488 (CONC).
89
Gayana Bot. 71(1), 2014
FIGURE 11. Pseudognaphalium landbeckii. A, Habit; B, Leaf pubescence; C, D, Foliar trichomes; E, Capitulum; F, Phyllaries; G, Pistillate
floret; H, Bisexual floret. I, Distribution [A, B, E, G, H, Landbeck in XII-1861, LP; C, D, F, Cabrera 11462, LP].
FIGURA 11. Pseudognaphalium landbeckii. A, Planta; B, Pubescencia de la hoja; C, D, Tricomas foliares; E, Capítulo; F, Filarios; G, Flor
pistilada; H, Flor perfecta; I, Distribución [A, B, E, G, H, Landbeck en XII-1861, LP; C, D, F, Cabrera 11462, LP].
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Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
8. Pseudognaphalium munoziae N. Bayón, C. Monti & S.E.
Freire, Phytotaxa 105(1): 1-10. 2013. TYPE: Chile: Región
Arica-Parinacota: Subida a Portezuelo de Chapiquiña,
18º20´1.7˝S, 69º31´1˝W, 4.147 m, 5-V-2011, A. Moreira,
M. Muñoz & V. Morales 1676 (holotype SGO!). Fig.12.
Perennial herbs, 4-6(15) cm tall, multistemmed, stems erect
or ascending, unbranched, arachnoid-glandulose. Stem
leaves remote, 10–25(30–40) × 1(2) mm, linear, margin
flat, apex acute to subobtuse, base attenuate; basal leaves
rosulate, 20–50(70–75) × 1-2 mm, linear, apex acute to
subobtuse, base long-attenuate; concolorous, greenish,
arachnoid-glandular on both surfaces, trichomes of 2 types,
long eglandular eseptate and long (short) glandular biseriate
trichomes. Capitula numerous, in clusters arranged in narrow
panicles, clusters subtended by linear leaves. Involucre 5–6
× 4–7 mm, campanulate. Phyllaries 3–4-seriate, lamina
opaque milky-white; outer phyllaries 4–5 × 1–2 mm,
narrowly ovate, apex obtuse; inner phyllaries 5–6 × 0.7–1
mm, ovate-oblong, apex obtuse. Pistillate florets 41–45;
corollas whitish, filiform, ca. 3 mm long. Bisexual florets
ca. 6, corollas yellowish at apex, tubular, 3.5–3.8 mm long.
Achenes ca. 0.3 mm long, glabrous, epidermis papillose
above. Pappus bristles 3.5–4 mm long.
PHENOLOGY. Flowering in May, June and October.
DISTRIBUTION AND HABITAT. Endemic to northern Chile (Arica
y Parinacota, Tarapacá), between 3500-4200 m.
RELATIONSHIPS. Pseudognaphalium munoziae is most similar
to P. psilophyllum. Both have campanulate involucres and
glandular linear leaves. Pseudognaphalium munoziae is
recognized by its conspicuously rosulate basal leaves and
shorter upper leaves, all apically acute to subobtuse, and
capitula arranged in narrow panicles with milky-white
phyllaries. Pseudognaphalium psilophyllum is distinguished
by its similar basal and upper leaves, which are attenuate
and acute at the apex, and capitula arranged in corymbs or
corymbose-paniculate with usually brown phyllaries
ADDITIONAL MATERIAL EXAMINED. CHILE. Región de
Arica y Parinacota. Prov. Parinacota, inicio de bajada a
Quebrada Allane, 18º0´7˝S, 69º37´14˝W, 3550 m, 10-VI2012, Moreira 1953 (SGO). Región de Tarapacá. Prov. El
Tamarugal, trayecto entre Enquelga y Berenguela, 9-X1997, Villagrán et al. 9229 (CONC).
9. Pseudognaphalium psilophyllum (Meyen & Walp.)
Anderb., Opera Bot. 104: 147. 1991. Gnaphalium
psilophyllum Meyen & Walp., Nov. Actorum Acad. Caes.
Leop. Carol. Nat. Cur. 19 (Suppl. 1): 275. 1843. TYPE:
Chile. Región de Valparaíso. Prov. Marga Marga, Limache,
9-XI-1927, 900 m s.m., Garaventa 1151 (neotype, designated
here, CONC!). Fig. 13.
Gnaphalium glandulosum Klatt, Linnaea 42: 129. 1878.
Pseudognaphalium glandulosum (Klatt) Anderb., Opera
Bot. 104: 147. 1991, syn. nov. TYPE: Chile. Parinacota,
Tacora, F.J.F Meyen s.n. (holotype, B (probably destroyed),
= photo F15099!; lectotype, here designated, GH 00008359,
= photo! fragment ex Herb. Klatt).
Perennial herbs, (5)10–85 cm tall, multistemmed, rarely
solitary, erect or ascending, usually branched at the upper
part, arachnoid-glandular. Stem leaves (7)10–65 × 1–6
mm, stem leaves linear, margin flat and usually revolute,
apex long-attenuate, acute, base not broad, decurrent 4–10
mm long; basal leaves 85–90 ×4–8 mm, linear-obovate;
concolorous, glandular on both surfaces, sometimes
arachnoid on the abaxial surface, trichomes of 2 types, long
(short) biseriate glandular trichomes and few long eglandular
eseptate, 3–6-celular trichomes. Capitula numerous in
clusters arranged in corymbs or panicles. Involucre broadly
campanulate, 3–5 × 3–5 mm. Phyllaries 4-seriate, lamina
straw-yellow to brown, hyaline and shiny, apex semiobtuse;
outer phyllaries 4.5–5 × 1.8–2.6 mm; inner phyllaries 3.7–
4.5 × 0.6–1.7 mm, oblong-obovate, apex acute. Pistillate
florets (46)82–143, corolla yellow, filiform, 2.5-3 mm long.
Bisexual florets 5–14, corolla yellow, tubular, 3–3.2 mm
long. Achenes 0.6–1 mm long, glabrous, epidermis smooth
to papillose. Pappus bristles 2.5–3.4 mm long.
PHENOLOGY. Flowering from September to May.
DISTRIBUTION AND HABITAT. Bolivia (Hind 2011), Argentina
(Jujuy, Salta, Catamarca, La Rioja, Mendoza, Neuquén,
Río Negro) and Chile (Arica y Parinacota, Tarapacá,
Antofagasta, Coquimbo, Valparaíso, Libertador General
Bernardo O´Higgins, Maule, Los Lagos), growing between
(5–50) 900–4500 m.
RELATIONSHIPS. Pseudognaphalium psilophyllum resembles
to P. munoziae. Both species are glandular or glandulararachnoid plants and have linear leaves. Pseudognaphalium
psilophyllum can be distinguished by its similar basal and
upper leaves (vs. conspicuously rosulate basal leaves and
shorter upper leaves in P. munoziae) and capitula in clusters
arranged in corymbs or corymbose-paniculate with usually
brown phyllaries (vs. capitula arranged in narrow panicles
with milky-white phyllaries).
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FIGURE 12. Pseudognaphalium munoziae. A, Habit; B, Leaf pubescence; C, D, Foliar trichomes; E, Capitulum; F, Phyllaries; G, Pistillate
floret; H, Bisexual floret. I, Distribution [A–H, from Freire et al. 2013].
FIGURA 12. Pseudognaphalium munoziae. A, Planta; B, Pubescencia de la hoja; C, D, Tricomas foliares; E, Capítulo; F, Filarios; G, Flor
pistilada; H, Flor perfecta; I, Distribución [A–H, tomado de Freire et al. 2013].
92
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
FIGURE 13. Pseudognaphalium psilophyllum. A, Apical flowering stem; B, Leaf ; C, D, Foliar trichomes; E, Capitulum; F, Phyllaries;
G, Pistillate floret; H, Bisexual floret without pappus; I, Distribution [A, B, E, G, H from Cabrera 1971; C, D, Spegazzini s.n., BAB; F,
Niemeyer et al. 8970, CONC].
FIGURA 13. Pseudognaphalium psilophyllum. A, Parte superior de la planta; B, Hoja; C, D, Tricomas foliares; E, Capítulo; F, Filarios;
G, Flor pistilada; H, Flor perfecta sin el papus; I, Distribución [A, B, E, G, H tomado de Cabrera 1971; C, D, Spegazzini s.n., BAB; F,
Niemeyer et al. 8970, CONC].
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Gayana Bot. 71(1), 2014
OBSERVATIONS
1. The types designated in the protologue of Gnaphalium
psilophyllum Meyen & Walp., ‘(Besser!) cum praecedente
(Meyen!)’, have not been located at B (probably destroyed),
GOET, M, or W, no illustration was published with the
name, and no other original material has been found. The
specimen ‘Garaventa 1151’ kept at CONC, which is in
accordance with the protologue and matches the locality
(‘cum praecedente’, i.e. Valparaiso in Gnaphalium dysodes
Spreng.) indicated in the protologue, is here selected as the
neotype of G. psilophyllum.
2. Klatt (1878) described Gnaphalium glandulosum on
the basis of its ‛caule fruticoso ascendente erecto ramoso
follisque glandulosis, foliis lanceolato-linearibus acuminatis
semiamplexicaulibus,
ramis
paniculato-corymbosis,
capitulis conglomeratis, involucri campanulati basi lanati
squamis obtusis pallide-fuscescentibus’. Gnaphalium
glandulosum is proposed here as a new synonym of G.
psilophyllum because there is no diagnostic character that
differentiates the former from the latter. Both have erect
or ascending stems that are arachnoid-glandular, caulinar
leaves which are linear, attenuate at the apex, decurrent
at the base, concolorous, glandular on both surfaces, and
capitula in clusters arranged in corymbs or panicles.
ADDITIONAL MATERIAL EXAMINED. CHILE. Región de Arica y
Parinacota. Prov. Arica, camino de Arica al Portezuelo de
Chapiquiña, km 93, 3350 m, 8-II-1964, C. Marticorena et al.
21 (CONC); camino de Azapa a Chapiquiña, km 76, 3250
m, 24-IX-1958, Ricardi & C. Marticorena 4730 (CONC);
FF.CC. de Arica-La Paz, Puquios, km 106-108, 3600-3700
m, 27-IX-1955, Ricardi & Silva 3535 (CONC, LP). Prov.
Parinacota, Socoroma, 3200 m, 2-II-2000, Belmonte 20060
(CONC); Putre, a 4 km, 18º12´S - 69º34´W, 3500 m, 16-V1989, Niemeyer et al. 8970 (CONC); Putre, 3500 m, 12II-1964, C. Marticorena et al. 141 (CONC); Putre, about
4-6 km below turnoff to Putre on road to Arica, ca. 3500
m, 29-XII-1995, Landrum & Landrum 8887 (MO); camino
a Chapiquiña, quebrada frente a la central hidroeléctrica,
3400 m, 8-V-1972, Ricardi et al. 286 (CONC); cordillera
de Arica, camino de Parinacota a Lago Chungará, 4200 m,
29-III-1961, Ricardi et al. 311 (CONC); entre Zapahuira
y Putre, 3270-3600 m, 15-IV-1980, Villagrán et al. 2441
(CONC); camino de Arica a Zapahuira, 2900 m, 18°28’S,
69°45’W, 13-IV-1980, Villagrán et al. 2353 (CONC); entre
Zapahuira y Putre, 3270-3600 m, 18º18’S, 69º34’W, 15-IV1980, Villagrán et al. 2478 (CONC); Socoroma, 3200 m,
12-VII-1982, Zöllner 11254 (MO). Región de Tarapacá.
Prov. El Tamarugal, camino de Huara a Cancosa, cerca de
Coscaya, 3200 m, 2-IV-1961, Ricardi et al. 398 (CONC);
trayecto entre Enquelga y Berenguela, 4200 m, 19º15’S,
69º08’W, 9-IX-1997, Villagrán et al. 9237 (CONC);
Prov. Iquique, Quebrada de Chusmiza, 19º41´S-69º11´W,
94
4-IX-1947, Villagrán et al. 9098 (CONC); Cordillera
Cerro Japu, 4000 m, III-1926, Wedermann 1100 (CONC,
LP, SI); Interior de las Termas de Mamiña, 3200 m, 4-II1965, Zőllner 976 (LP). Región de Antofagasta. Prov. El
Loa, Quebrada Del Inca, Cerro Aucanquilcha, 4500 m,
21°14’S, 68°28’W, 2-IV-1985, Arroyo 85-586 (CONC);
camino de San Pedro de Atacama a Paso Jama, límite entre
pastizal y tolar, 4200 m, 4-IV-1997, Arroyo et al. 97069
(CONC), transecto Cerro Carasilla a Sala Ascotán, 4350
m, 28-III-1985, 21°41’S, 68°17’W, Arroyo et al. 85-393
(CONC). Región de Coquimbo. Prov. Choapa, in Potrero
Largo near Cuncumén, 2700 m, 6-II-1984, Zöllner 11991
(MO); Alrededores de Toconce, terrazas de cultivo, 3300
m, 22°15’S, 68°10’W, 26-I-1980, Villagrán & Armesto
2271 (CONC); Illapel, Palo Colorado, pleno campo, 10 m,
32°04’S, 71°31’W, 10-XI-1961, Jiles 3887 (CONC); Prov.
Elqui, Carretera Panamericana entre Pichidangui y Los
Vilos, 12-X-1963, C. Marticorena & Matthei 49 (CONC);
ibid., C. Marticorena & Matthei 67 (CONC). Región de
Valparaíso. Prov. Valparaíso, Algarrobo, 20 m, XI-1965,
Semler s.n., Herb. Gunckel 48696 (CONC 116219);
Tunquén, estero Casablanca, 10-50 m, 33°16’S, 71°39’W,
1-XI-2001, Teillier 5915 (CONC). Región del Libertador
General Bernardo O’Higgins. Prov. Cachapoal, valle
superior Río Cachapoal, 1500 m, X-1978, Prov. Colchagua,
Termas de El Flaco, 23-I-1948, Barros 7332 (LP). Región
del Maule. Prov. Curicó, Cord. del Planchón, 17-II-1939,
Barros 2303 (SI). Región de Los Lagos. Prov. Chiloé, isla
Surgidero, 43º25´S - 74º14´W, 5-30 m, 9-II-1986, Villagrán
& Leiva 7448 (CONC).
10. Pseudognaphalium remyanum (Phil.) Anderb., Opera
Bot. 104: 147. 1991. Gnaphalium remyanum Phil., Anales
Univ. Chile 90: 8. 1895. TYPE: Chile. Santiago, XII-1877,
Philippi s.n. (lectotype, here designated SGO 44975!;
isolectotypes, B (probably destroyed), = photo F15113!,
SGO 64436!). Fig. 14.
Perennial herbs, ca. 50 cm tall, stem branched above
base, whitish-arachnoid. Leaves up to 50 × 8 mm, linear,
margin flat, apex acuminate, base decurrent 3–4 mm
long, and sometimes broad; discolorous, adaxial surface
green, abaxial surface whitish, arachnoid on both surfaces,
trichomes of 2 types, long eglandular eseptate, 3-celular,
and few short biseriate glandular trichomes. Capitula
numerous in clusters arranged in corymbs or lax panicles.
Involucre broadly campanulate, 2.2–2.5 × 2–4 mm.
Phyllaries 3–4-seriate, lamina yellowish to whitish, hyaline
and shiny; outer phyllaries ca. 2 × 1.6 mm, broadly ovate,
apex rounded; inner phyllaries ca. 2 × 0.8 mm, oblongobovate, apex obtuse. Pistillate florets ca. 163, corolla
yellow, filiform, 1.2 mm long. Bisexual florets ca. 13,
corolla yellow, tubular, 1.5 mm long. Achenes ca. 0.3 mm
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
FIGURE 14. Pseudognaphalium remyanum. A, Habit; B, Leaf pubescence; C, D, Foliar trichomes; E, Capitulum; F, Phyllaries; G, Pistillate
floret; H, Bisexual floret. I, Distribution [A, E–H, Philippi in 1877, SGO; B–D, Buchtien 174, NY].
FIGURA 14. Pseudognaphalium remyanum. A, Planta; B, Pubescencia de la hoja; C, D, Tricomas foliares; E, Capítulo; F, Filarios; G, Flor
pistilada; H, Flor perfecta; I, Distribución [A, E–H, Philippi en 1877, SGO; B–D, Buchtien 174, NY].
95
Gayana Bot. 71(1), 2014
long, glabrous, epidermis papillose. Pappus bristles 1.3–1.7
mm long.
Tucumán) and northern-central Chile (Arica y Parinacota,
Tarapacá, Antofagasta, Coquimbo, Metropolitana, Maule,
Biobío), growing between (20/300-800) 2600-3500 m.
PHENOLOGY. Flowering in December.
DISTRIBUTION AND HABITAT. Bolivia and Central Chile (Región
Metropolitana).
RELATIONSHIPS. Pseudognaphalium remyanum seems to be
distinguished by its whitish arachnoid plants, linear leaves,
and capitula in clusters arranged in lax corymbs or panicles
with broadly ovate outer phyllaries.
OBSERVATION
Although no Chilean material was available of
Pseudognaphalium remyanum, it closely resembles the
genus Pseudognaphalium in habit, leaves, floral characters
and arrange of its capitula, so the species is referred here,
until further study.
ADDITIONAL MATERIAL EXAMINED. BOLIVIA. Sur-Yungas,
Sirupaya vec. de Yanacachi, 14-XII-1906, O. Buchtien 174
(NY).
11. Pseudognaphalium tarapacanum (Phil.) Anderb.,
Opera Bot. 104: 148. 1991. Gnaphalium tarapacanum
Phil., Anales Mus. Nac., Santiago de Chile 8: 46. 1891.
TYPE: Chile. Tarapacá, Paroma 25-II-1885, F. Philippi s.n.,
Calcahuay, I-1886, Rahmer s.n. (lectotype, here designated,
SGO 64433! - entire plant). Fig. 15.
Perennial herbs, 10–20 cm tall, multistemmed, stems
erect or ascending, unbranched or branched at the upper
part, arachnoid-glandulose. Stem leaves ca. 40 × 4–6
mm, lanceolate, margin flat, apex attenuate, base broad
and decurrent 5–7 mm long; basal leaves oblanceolatespathulate, apex acute or subotuse; concolorous, arachnoidglandulose on both surfaces, trichomes of 2 types, long
eseptate eglandular trichomes, 5-8 celular and few short
biseriate glandular trichomes. Capitula numerous arranged
in dense terminal clusters. Involucre broadly campanulate,
3–5 × 3–4 mm. Phyllaries 3–4-seriate, lamina light brown,
hyaline and shiny, apex rounded; outer phyllaries 3–5 ×
(0.5)1.5–2 mm, elliptic; inner phyllaries 3–4 × 0.5–1 mm,
oblong. Pistillate florets 66–151, corolla yellowish, filiform,
2–3 mm. Bisexual florets 5–11, corolla yellowish, tubular,
2–3 mm. Achenes 0.5–0.9 mm long., pilose, with short
duplex hairs or glabrous with epidermis papillose. Pappus
bristles, 2–3 mm long.
PHENOLOGY. Flowering from (November) January to May.
DISTRIBUTION AND HABITAT. Northwestern Argentina
(Catamarca, Córdoba, Jujuy, La Rioja, Salta, San Luis,
96
RELATIONSHIPS. Pseudognaphalium tarapacanum resembles
P. psilophyllum. Both species are glandular or glandulararachnoid plants. Pseudognaphalium tarapacanum is
easily distinguished by its stem leaves broad at base and
occasionally undulate leaf blade margins base (vs. stem
leaves not broad at base and flat leaf blade margins in P.
psilophyllum).
OBSERVATION
The protologue of Gnaphalium tarapacanum mentions two
specimens from Tarapacá Province, i.e. ‘Habitat in provincia
Tarapacá ad Paroma et Calcahuay’. We located at LP the
collection Philippi s.n., Paroma, 25 II 1885 LP 001945, at
SGO one sheet-SGO 64433 with two specimens (an entire
plant, on the right hand side and a fragment, on the left
hand side) mounted on it, and two labels on the right-hand
side, i.e. ‘Calcahuay, Januario 1886, Rahmer’ and ‘Paroma,
25 II 1885, F. Ph’, and a photo F15120 ‛Tarapacá, Phil.’.
We propose the specimen kept at SGO as the lectotype
where the original herbarium of Philippi is kept, and which
represent the most complete plant. Since the collector and
collection number can not be assigned with certainty to each
mounted specimen, we selected as lectotype of Gnaphalium
tarapacanum the plant on the right hand of the sheet.
ADDITIONAL MATERIAL EXAMINED. CHILE. Región de Arica y
Parinacota. Prov. Arica, Lago Chungará, ribera occidental,
18º14´S, 69º10´W, 4500 m, 20-V-1979, Villagrán et al. 1271
(CONC); camino de Arica al Portezuelo de Chapiquiña, km
93, 3350 m, 8-II-1964, C. Marticorena et al. 29 (CONC);
camino a Chapiquiña, Quebrada frente a la Central
Hidroeléctrica, 3400 m, 8-V-1972, Ricardi et al. 279 (CONC);
Prov. Parinacota, Putre, 3500 m, 23-XI-1955, Levi Heins 306
(CONC). Región de Tarapacá. Prov. El Tamarugal, camino
de Cancosa al límite con Bolivia, km 2, 4000 m, 18-II-1964,
Marticorena et al. 360 (CONC); 2 km al E de Chuzmisa,
3520 m, 19°40´S-69°09´W, 2-IV-2007, Rosas 4946 (CONC).
Región de Antofagasta. Prov. El Loa, Chiuchiu, en una huerta,
2560 m, 3-I-1950, Pfister s.n. (CONC 9364). Región de
Coquimbo. Prov. Elqui, camino entre Embalse La Laguna y
Campamento del Embalse, 30º12´S, 70º02´W, 2900-3200 m,
8-I-1981, Arroyo 81112-A (CONC). Región Metropolitana de
Santiago. Prov. Santiago, Cerro San Cristóbal, 800 m, 14-V1954, Navas 678 (CONC). Región del Maule. Prov. Linares,
a lo largo del río Achibueno, desde junta estero Las Ánimas
a junta estero Riecillo, 30º05´S, 71º10´W, 9-III-1999, Ruiz
& López 1231 (CONC). Región del Biobío. Prov Arauco,
Isla Mocha, Laguna Hermosa, 300 m, 15-III-1971, Weldt &
Rodríguez 1149 (CONC); Prov. Ñuble, Cobquecura, 20 m,
13-IV-1954, Ledezma 225 (CONC).
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
FIGURE 15. Pseudognaphalium tarapacanum. A, Habit; B, Leaf base; C, Leaf pubescence; D, E, Foliar trichomes; F, Capitulum; G,
Phyllaries; H, Distribution [A, B, F, from Cabrera 1978; C-E, Marticorena et al. 360, CONC; G, Marticorena et al. 29, CONC].
FIGURA 15. Pseudognaphalium tarapacanum. A, Planta; B, Base de la hoja; C, Pubescencia de la hoja; D, E, Tricomas foliares; F, Capítulo;
G, Filarios; H, Distribución [A, B, F, tomado de Cabrera 1978; C-E, Marticorena et al. 360, CONC; G, Marticorena et al. 29, CONC].
97
Gayana Bot. 71(1), 2014
12. Pseudognaphalium viravira (Molina) Anderb., Opera
Bot. 104: 148. 1991. Gnaphalium viravira Molina, Sag.
Stor. Nat. Chili 149, 354. 1782. TYPE: Chile. Valparaíso
‛La Vinna de la mar’, IX-1830, C. L. G. Bertero 1823
(lectotype, here designated, CONC!; isolectotype, NY
00169526,= photo!). Fig. 16.
Gnaphalium montevidense Spreng., Syst. Veg. 3: 475. 1823.
Pseudognaphalium montevidense (Spreng.) Anderb. Opera
Bot. 104: 147. 1991, syn. nov. TYPE: Uruguay. Montevideo,
Sello s.n. Herb. Sprengel 785 (holotype, P 704564, = photo!;
isotipo, LP 001932! fragment ex P).
Gnaphalium coquimbense Phil., Linnaea 29: 5. 1858.
Pseudognaphalium coquimbense (Phil.) Anderb., Opera
Bot. 104: 147. 1991, syn. nov. TYPE: Chile. Coquimbo,
La Serena, IX-1837, C. Gay 678 (holotype: SGO 64422!;
isotype, B (probably destroyed ), = photo F15094 SI!).
Gnaphalium pratense Phil., Linnaea 33: 166. 1864.
Pseudognaphalium pratense (Phil.) Anderb., Opera Bot.
104: 147. 1991, syn. nov. TYPE: Chile. Valdivia, Ranco,
I-1860, R.A. Philippi s.n. (lectotype designated by Cabrera
(1971: 112), SGO 71286!).
Gnaphalium illapelinum Phil., Linnaea 33: 164. 1864.
Pseudognaphalium illapelinum (Phil.) Anderb., Opera Bot.
104: 147. 1991, syn. nov. TYPE: CHILE. Coquimbo, Illapel,
XII-1862, L. Landbeck s.n. (lectotype, here designated SGO
64423!; isolectotypes, B (probably destroyed), = photo
F15101!, LP 001923!, NY!, SGO 44966!).
Gnaphalium andicola Phil., Anales Univ. Chile 90: 17.
1895. Pseudognaphalium andicola (Phil.) C. Monti, N.
Bayón & S.E. Freire, J. Bot. Res. Inst. Texas 7(1): 196.
2013, syn. nov. TYPE: Chile. Santiago, Las Condes, L.
Navarro s.n. (lectotype, designated by Monti et al. (2013:
196), SGO 64481!).
Gnaphalium longifolium Phil., Anales Univ. Chile 90: 13.
1895, syn. nov. TYPE: Chile. Valdivia, S. Juan, II-1882,
Philippi s.n. (holotype, SGO 64446!; isotype, LP 001930!).
Gnaphalium subnudum Phil., Anales Univ. Chile 90: 15.
1895, syn. nov. TYPE: Chile. ‛Habitat in andibus sed quo
loco?’ without collector (lectotype, here designated, SGO
64429!).
Perennial herbs, 5–40(60) cm tall, multistemmed, stems erect
or ascending, unbrached, more rarely branched at the upper
part, whitish-woolly. Stem leaves 13-50 (60) × 1–5(10)
mm, linear or linear-ovate to linear-oblong, margin flat or
occasionally undulate, apex acute to attenuate, base clasping
and decurrent 2–7(10) mm long; basal leaves commonly
98
approximate, 23–50 × 3–6(10) mm, linear-obovate, apex
obtuse, base long-attenuate; concolorous, whitish-woolly
to arachnoid-glandulose on both surfaces, trichomes of 2
types, long eseptate eglandular, 3–6-celular and very few
short biseriate glandular trichomes hidden under the wool.
Capitula numerous in dense terminal clusters, sometimes
arrranged in corymbs. Involucre campanulate, 3–5 × 3–5
mm. Phyllaries 3−4-seriate, lamina yellowish or light
brown to whitish, hyaline and shiny; outer phyllaries 2.8–
4.2 × 1.4–2 mm, obovate, apex obtuse or acute; inner
phyllaries 3–4.3 × 1–1.1 mm, linear-elliptic, apex acute
or subobtuse. Pistillate florets, (82–88)115–130, corolla
yellow, filiform, (2.8)3.7–4 mm long. Bisexual florets 5–16,
corolla yellow, tubular, (2.7)3.3–3.7 mm long. Achenes 0.4–
0.5 mm long, glabrous with smooth or papillose epidermis
or pilose with short oblong duplex hairs, sometimes with
papillose epidermis and few short oblong duplex hairs.
Pappus bristles (2.2)3.2–4 mm long.
Pseudognaphalium viravira is treated here as a polymorphic
species, with plants ranging from few centimetres to near
half meter tall, upper leaves varying from linear to linearobovate, acute to attenuate at the apex, with margins flat
or rarely obscurely undulate, usually woolly to more rarely
arachnoid-glandulose, and glabrous to pilose achenes.
PHENOLOGY. Flowering from August to March.
VERNACULAR NAME. ‛Nido de diuca’, ‛vira-vira’.
DISTRIBUTION AND HABITAT. Chile (Tarapacá, Atacama,
Coquimbo, Valparaíso, Metropolitana, Libertador General
Bernardo O´Higgins, Maule, Biobío, Araucanía, Los Lagos,
Aysén del General Carlos Ibáñez del Campo, and probably
Magallanes y de la Antártica Chilena) and central-southern
Argentina (Buenos Aires, Chubut, Jujuy, La Rioja, Mendoza,
Neuquén, Río Negro, Salta, San Juan, Santa Cruz, Tierra del
Fuego), growing on dry soils or in forest in shady places;
from sea level to 3200 m.
RELATIONSHIPS. Pseudognaphalium viravira can be
distinguished by its woolly mutistemmed plants, unbranched
stems, basal leaves usually approximate, upper leaves usually
remote, and capitula in dense terminal clusters (arranged in
corymbs when plants fully developed). Pseudognaphalium
viravira is closely related to the cosmopolitan species
Pseudognaphalium luteoalbum (L.) Hilliard & B.L.
Burtt. Both species have woolly leaves on both surfaces;
approximate basal leaves with linear-spathulate, apically
obtuse leaf blade; remote stem leaves with oblong-linear,
apically acute, occasionally undulate margin leaf blade, and
capitula arranged in dense terminal clusters with involucre
3–5 mm high. P. luteoalbum differs from this species by its
annual plants (vs. perennial in P. viravira) and yellowish
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
FIGURE 16. Pseudognaphalium viravira. A, Habit; B, Leaf pubescence; C, D, Foliar trichomes; E, Capitulum; F, Phyllaries; G, Pistillate
floret; H, Bisexual floret. I, Distribution [A–H, Gunckel 39466, CONC].
FIGURA 16. Pseudognaphalium viravira. A, Planta; B, Pubescencia de la hoja; C, D, Tricomas foliares; E, Capítulo; F, Filarios; G, Flor
pistilada; H, Flor perfecta; I, Distribución [A–H, Gunckel 39466, CONC].
99
Gayana Bot. 71(1), 2014
florets, usually reddish at apex (vs. whitish in P. viravira).
Although P. luteoalbum was mentioned to RME (Rémy 1849,
Iboda 2014), we have not seen specimens of this species.
OBSERVATIONS
1. The protologue of Gnaphalium viravira did not include
type collection. Later, Candolle (1838: 224), mentioned for
G. viravira, two specimens collected by Bertero, i.e. ‘Bert.
n. 1032 et 1823’ from Quillota and Valparaíso, respectively.
We located the specimen ‘Bertero1823’ at CONC, and at
NY a sheet with two specimens mounted on it, one with
double collection numbers Bertero 309 & 1032 (NY
00169525),‛In pascuis petrosis calidis Quillota, et ad sepes
secus vias prope La Quinta, Chili. Octbr. Nvbr. 1829’, and
the other Bertero 1823 (NY 169526), ‛la Vinna de la mar.
Valparaiso. Chili. Spbr. 1830’. A third sheet of Bertero 309
& 1032 was located at MO (MO 5448098, = photo!). We
propose to select as lectotype the specimen ‘Bertero1823’
kept at CONC which presents the most complete plant, and
to exclude from type material the specimen Bertero 309 &
1032 (NY 169525, MO 5448098) because of its ambiguous
information.
2. After examining the type materials, Gnaphalium
montevidense, G. pratense, and G. andicola are proposed
here as new synonyms of G. viravira because there is no
diagnostic character to separate them as distinct taxa.
All of these are multistemmed whitish-woolly plants,
with approximate basal leaves and remote stem leaves
that are linear to linear-obovate and apically acute, and
capitula arranged in dense terminal clusters. In addition,
the differential character mentioned in the protologue of
Gnaphalium montevidense, i.e., stem leaves clasping, is a
variable feature throughout the range of G. viravira.
3. Gnaphalium andicola was placed into synonymy
of Pseudognaphalium cheiranthifolium (Hind 2011).
According to the protologue of G. andicola and Gnaphalium
cheiranthifolium, they have leaves ‛lanceolato-linearibus
longe acuminatis’ and ‛foliis lineari-lanceolatis acutis’,
respectively. However, examination of the original material
of both species at SGO and P, respectively reveals that the
leaves of Gnaphalium andicola are linear to narrowly linearobovate, apically acute (vs. lancelolate to linear-lanceolate,
apically attenuate in G. cheiranthifolium). In addition,
Gnaphalium andicola differs from P. cheitanthifolium by
it multistemmed plants (vs. solitary stems, rarely 2 or 3,
in P. cheiranthifolium), and its capitula usually in solitary
terminal clusters (vs. capitula in cluster arranged in coryms
in P. cheiranthifolium).
4. After examining the type materials, Gnaphalium
coquimbense is proposed here as a new synonym of
G. viravira because there is no diagnostic character
100
that differentiates the former from the latter. Both are
multistemmed whitish-woolly plants, with linear spathulate
basal leaves and remote stem leaves that are apically
acuminate, shortly decurrent at the base. In addition, the
differential characters mentioned in the protologue of G.
coquimbense, i.e., congested inflorescences and yellowish
phyllaries, are variable features throughout the range of G.
viravira.
5. The protologue of Gnaphalium illapelinum mentions
the collection ‘Prope Illapel, Lanbeck lectum’. We located
at SGO and LP the collection ‘Illapel, December 1862’,
SGO 44966, 64423; LP 001923, and the colection ‘Illapel,
prov. Coquimbo’ at NY. Even if these four specimens are in
accordance with the protologue, we selected as the lectotype
of Gnaphalium illapelinum the specimen SGO 64423,
which presents the most complete plant and has Cabrera’s
annotation ‘Typus’.
6. After examining the type materials, Gnaphalium
illapelinum is proposed here as a new synonym of G. viravira
because there is no diagnostic character that differentiates
the former from the latter. Both are multistemmed whitishwoolly plants, with approximate basal leaves and remote
stem leaves that are apically attenuate, and capitula arranged
in dense terminal clusters. In addition, the differential
character mentioned in the protologue of G. illapelinum,
i.e., arachnoid glandulose leaves, is a variable feature
throughout the range of G.viravira.
7. According to the protologue, Gnaphalium subnudum was
based on three specimens without locality, i.e. ‘tenemos
tres ejemplares, de los cuales el mayor …’. We found at
SGO a sheet with two specimens mounted on it, which has
Philippi’s annotation on the right-hand side ‘Gnaphalium
subnudum’. This specimen which is in accordance with the
protologue of Gnaphalium subnudum, is here designated as
the lectotype.
8. Gnaphalium longifolium Phil. and G. subnudum Phil.,
which were previously placed into synonymy of G. pratense
Phil. and G. montevidense, respectively (Cabrera 1971),
are here considered as synonyms of Pseudognaphalium
viravira.
ADDITIONAL MATERIAL EXAMINED. CHILE. Región de
Tarapacá. Prov. El Tamarugal, Termas de Mamiña, 2-4 km
antes del pueblo, 2700 m, 18-X-1965, Ricardi et al. 1348
(CONC); Mamiña, 2750 m, IX-1956, Richter s.n. (CONC
116230). Región de Atacama. Prov. Huasco, Río Laguna
Grande, entre Quebrada Candelilla y el extremo oeste de la
Laguna Grande, 2800-3100 m, 2-I-1983, C. Marticorena et
al. 83413 (CONC); S/Prov., Estancia Manflas, 1450 m,
2-XI-1956, Ricardi & C. Marticorena 3747 (CONC).
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
Región de Coquimbo. Prov. Choapa, Illapel, 19-IX-1926,
Barros 1466 (CONC); ídem, 310 m, VIII-1971, Zöllner
5133 (CONC); ídem, s/fecha, Philippi s.n. (NY); Los Vilos
near the coast, 30-X-1976, Zöllner 9914 (CONC, MO);
Valle Ojotas, Potrero Largo, 3200 m, 8-II-1984, Zöllner
12142 (SI); Cordillera de Combarbalá, Potrero Grande,
Ramadilla, 31º18S, 70º50W, 2600 m, 6-I-1963, Jiles 4428
(CONC); Quilimari, 5 m, 9-IX-1982, Montero 12281
(CONC); Prov. Elqui, entre La Serena y la Cuesta de Las
Chilcas, orillas de la Carretera, 20-IX-1974, Mahu 10213
(LP); Punta de Teatinos, II-1950, Chang 20965 (CONC); La
Serena, 15 m, IX-1926, Barros 2424 (CONC); ídem, I-1928,
Barros 21755 (CONC); ídem, IX-1929, Barros 2840
(CONC); Valle del Elqui, El Molle, road leading E from
village, along the N side of the river, 29º58’51’’S,
70º56’17’’W, 365 m, 7-XI-2006, Tepe et al. 1840 (CONC);
Coquimbo, besides the Panamerican near Teniente Bridge,
10-IX-1987, Zöllner 13652 (MO); Prov. Limarí, Ovalle, IX1926, Barros 2417 (CONC); Ovalle, Zorrilla, 30°50′S,
71°30′W, 350 m, 26-IX-1946, Jiles 910 (CONC); ídem, 17IX-1950, Jiles 1818 (CONC, LP); Ovalle, Parque Nacional
Fray Jorge, parte baja, 30°40′S, 71°33′W, 300 m, 5-XI1974, C. Marticorena et al. 421 (CONC); Ovalle, Carretera
Panamericana, 5 km al S de Socos, 28-X-1965, Ricardi et
al. 1551 (CONC); Cordillera de Ovalle, Cerro Loica,
31º02’S, 72º42’W, 2000 m, 18-XII-1965, Jiles 4735
(CONC); Hurtado Valley, 16-IX-1986, Zöllner 13111 (MO).
Región de Valparaíso. Aconcagua, Pichidangui, 470 m, 18II-1962, Mahu s.n. (LP); Prov. Marga Marga, Limache, 31V-1924, A.J.H. 680 (CONC 89681); ídem, 18-IX-1927,
Looser 203 (SI); Villa Alemana, 21-IX-1937, Barros 2301
(LP, SI); Valle de Marga-Marga, 40 km al E de Valparaíso,
IX-1916, Jaffuel 896 (GH); Marga-Marga, 150 m, IX-1910,
Jaffuel 899 (CONC); Prov. Quillota, Cuesta de Melón, 13IX-1957, Ricardi & C. Marticorena 4222 (CONC); Prov.
Petorca, Papudo, 28-X-1974, Zöllner 7936 (CONC); Rt. 5
between tunnel and La Ligua, 32º33’25’’S, 71º15’54’’W,
110 m, 3-XI-2006, Tepe et al. 1689 (CONC); Prov. San
Felipe de Aconcagua, Carretera Panamericana, Bajada de El
Melón, 13-IX-1957, Cabrera 12523 (LP); Prov. Valparaíso,
Viña del Mar, 24-IX-1921, Behn 20879 (CONC); Jardín
Botánico Nacional de Viña del Mar, 10-V-1956, Cabrera
12215 (LP); ídem, Sendero Marianne North, 33º2’15’’S,
71º29’38’’W, 21-I-2005, Novoa s.n. (CONC 165314);
Quintero, Los Molles, 32°46′S, 71°32′W, XII-1953,
Gunckel 40598 (CONC); Quilpué, 33°03’S, 71°27’W, 80
m, 13-XI-1969, Schlegel 3144 (CONC); Chorrillos, 26-XII1944, Boelcke 363 (SI); Concón, 15 m, XII-1951, Gunckel
23453 (CONC); Tranque Pitama, 250 m, 15-XI-2010,
Rabanales s.n. (CONC 173012). Región Metropolitana de
Santiago. Prov. Chacabuco, Batuco, 480 m, 25-IX-1951,
Gunckel 22760 (CONC); Prov. Cordillera, Clarillo, 800 m,
Gunckel 39466 (CONC); Prov. Maipo, Laguna de Aculeo,
350 m, 11-X-1942, Pisano et al. 1567 (CONC); Prov.
Santiago, Quebrada de Macul, XI-1965, Muñoz 48718
(CONC); Cerro de Renca, 750 m, XI-1950, Gunckel 22280
(CONC); Subida al Cerro Caléu, 33°00′S, 70°57′W, 800 m,
13-XI-1960, Schlegel 3125 (CONC); San José de Maipo,
Cajón de Morales, entre Panimávidas y la laguna, 2100 m,
25-I-2002, Teillier & Márquez 5315 (CONC); Monumento
Natural El Morado, Cajón del Morales, 33º49’S, 70º05’W,
2300 m, 29-XII-1990, Teillier & González 2301 (CONC,
MO); Cajón del Río Maipo, Quebrada El Canelo, 12-XII1973, Mahu 9795 (LP); Cajón del Maipú, Potrero Grande,
2800 m, 2-I-1967, Zöllner 1770 (LP); San José de Maipo,
Cajón del río Morales, 17-I-1989, Saavedra & Pauchard 111
(CONC); Alhué, Monte Santillana, 2-I-1939, Barros 2285
(LP, SI); Lo Valdés (Baños Colina), 2500 m, 2-I-1968,
Zöllner 3031 (CONC); Lo Valdés, 2300 m, 7-III-1953,
Ricardi 2448 (CONC); ídem, en las cercanías del Refugio
alemán, Quebrada Valdés, 2000 m, 20-XI-1940, Garaventa
5681 (CONC); entre Lo Valdés y las Yeseras, 2450 m, 11-II1963, Ricardi et al. 834 (CONC); Río Colorado, 2500 m,
I-1950, Moreno 25689 (CONC); mounts above Río
Colorado, 4000 ft, 21-I-1902, Hastings 418 (NY); Pirque,
Cerro Blanco (Puente Alto), 11-V-1970, Mahu 4885 (LP);
Quebrada de Ramón, 33º26’S, 70º 30’W, 1200-1600 m, 19XI-2000, Tomé 130 (CONC); Fundo Valencia, Monte
Águila, 1/3-XI-1939, Looser 3990 (LP); Prov. Talagante,
Peñaflor, 400 m, XI-1965, Arellano 48720 (CONC). Región
del Libertador General Bernardo O’Higgins. Prov.
Colchagua, San Fernando, 1700 m, 3-I-1959, Montero 6056
(CONC); San Fernando, Vegas Del Flaco, 1800 m, 7-II1955, Ricardi 3157 (CONC); Termas del Flaco, 34º56’S,
70º25’W, 1950 m, 3-II-1989, Niemeyer & Fernández 8908
(CONC); Río Tinguiririca, San Fernando, 300 m, 29-XII1950, Ricardi s.n. (CONC 9874); Prov. Cachapoal, Palmar
de Cocalán, 34º11’S, 71º14’W, 450 m, 28-I-1964, Schlegel
4949 (CONC). Región del Maule. Prov. Curicó, Cajón del
Lontué, I-1884, Philippi s.n. (sintipo de G. andicola, SGO
064385); Lolol, 17-IX-1937, Barros 184 (LP); 9 km al E de
Los Queñes, 35°01’32”S, 70°42’53”W, 815 m, 13-XI-2006,
Tepe 2003 (CONC); Cordillera del Potrero Grande, 30-I1928, Barros 12 (LP); Peteroa, I-1933, Grandjot s.n. (SI);
Peteroa, 2800 m, I-1933, C. & G. Grandjot s.n. (SI 141479);
Cordillera de El Planchón, 17-II-1939, Barros 2304 (LP,
SI); camino de Laguna de Teno a la junta con el camino
internacional a Paso Vergara, 2250 m, 10-III-1967, C.
Marticorena & Matthei 973 (CONC), 2300 m, C.
Marticorena & Matthei 967 (CONC); Prov. Linares,
Cauquenes, Fundo Porvenir, 250 m, 29-X-1975, Rodríguez
661 (CONC); Reserva Nacional Los Bellotos del Melado,
quebrada, 1415 m, 4-I-2000, Humaña et al. 20102 (CONC);
Reserva Nacional Los Bellotos del Melado, 1374 m,
Gardner et al. 295 (CONC); Altos de Vilches, camino a
Linares, fundo El Castillo, entrando por Copihue hacia el
este, 945 m, 6-III-1999, Ruiz & López 1018 (CONC); a lo
largo del río Achibueno, desde junta estero Las Ánimas a
101
Gayana Bot. 71(1), 2014
junta estero Riecillo, 36º05’S, 71º10’W, 9-III-1999, Ruiz &
López 1184 (CONC); Prov. Talca, Laguna del Maule, Talca,
2200 m, I-1943, Behn s.n. (CONC 4264, 20878); Maule,
8-II-1892, Kuntze s.n. (NY); Prov. Talca, Constitución,
Quivolgo, 26-XI-1958, A. Barnier 427 (CONC); Curepto,
I-1926, Barros 477 (CONC); camino de Talca hacia Putú,
fundo El Trapiche, 120 m, 4-I-1964, Matte 745 (CONC);
Altos de Vilches, camino a Laguna El Alto, 2000 m, 29-I2000, Finot & López 1804 (CONC); Laguna de la Invernada,
orilla oeste, 35°43´S-70°46´W, 1300 m, 11-IV-2000, Finot
& López s.n. (CONC); Comuna de San Clemente, Paso
Pehuenche, 1750 m, I-2005, Luebert & Teillier 2232
(CONC). Región del Biobío. Prov. Arauco, Playa Punta
Morguilla, 37º42’S, 73º38’W, 6-II-2001, Parra & Torres
431 (CONC); Prov. Biobío, Panamericana a Pangal del
Laja, km 10, 150 m, 37°11’S, 72°17’W, 29-XI-1976,
Marticorena & Rodríguez 8430 (CONC); Laguna de la
Laja, 21-I-1969, Cabrera 19677 (LP); Laguna del Laja,
faldeos Sierra Velluda, 15-II-1960, Ricardi & Marticorena
5150 (CONC); Laguna del Laja, Los Barros, 22-I-1969,
Ricardi & C. Marticorena 5766 (CONC); Laguna del Laja,
Los Barros, sector Mallín Florido, 1458 m, 23-I-2001,
Baeza et al. 3110 (CONC); faldeos del volcán Antuco,
frente a la Laguna del Laja, 21-I-1969, Ricardi & C.
Marticorena 5721 (CONC); Parque Nacional Laguna del
Laja, camino Los Barros a Pichachén, 1470 m, 23-I-2001,
Baeza et al. 3074 (CONC); entre Carretera Longitudinal y
Pangal del Laja, km 10, 37º11’S, 72º17’W, 29-XI-1976, C.
Marticorena & Rodríguez 8417 (CONC); El Roble, 2-II1929, Barros 22611 (CONC); Prov. Concepción, fundo
Bolslal, 5-XII-1933, Belisario 849 (CONC); camino a
Coronel, km 15, 7-XII-1962, Gleisner s.n. (CONC 89714);
camino de Rere a Yumbel, 6-I-1959, Marticorena et al. 43
(CONC); Pichaco, Hualqui, 12-VIII-1936, Junge s.n.
(CONC 5891); San Pedro, 1-XI-1943, Barros 3139 (SI);
Dunas de San Vicente, 24-X-1934, Junge 6285 (CONC);
Florida, Fundo Colico, 11-IX-1976, Oehrens s.n. (CONC
89742); camino de Hualqui a Rere, cerca de Gomero, 220
m, 5-I-1959, C. Marticorena et al. s.n. (CONC 25211); Prov.
Concepción/Ñuble, camino Concepción-Bulnes, 70 m,
6-XII-1944, Pfister s.n. (CONC 4949); Prov. Ñuble. Baños
de Chillán, 28-XII-1986, Zöllner 13147 (MO); Reserva
Nacional Ñuble, al Este de la avanzada de Cuatro Juntas,
junto al camino hacia la frontera, 1500 m, 7-II-2008, Teneb
551 (CONC); Reserva Nacional Ñuble, al Este de la
avanzada de Cuatro Juntas, junto al camino cerca de la pista
del gasoducto, 1500 m, 7-II-2008, Teneb 569 (CONC);
camino a termas de Chillán, Gruta Los Pangues, 1200 m,
7-XII-1992, Rodríguez & C. Marticorena 3053 (CONC);
Ñuble, Termas de Chillán, 1800 m, Cabrera 3636 (LP);
Termas de Chillán, 1750 m, 14-I-1967, Schlegel 5863
(CONC); Nevados de Chillán, Ruinas del Refugio Walldorf,
1940 m, 13-I-2009, Pfanzelt 173 (CONC). S/Prov., El
Roble, 3-II-1929, Barros 18 (LP); Chancho Co, 15-II-1947,
102
de Barba 2040 (LP); El Abanico, III-1949, Pfister s.n.
(CONC 8864). Región de la Araucanía. Prov. Cautín, Parque
Nacional Villarrica, 1070 m, 12-I-1993, Finckh 552
(CONC); Cunco, 13-I-1931, Barros 44 (LP); Laguna
Conguillío, 1150 m, 20-I-1976, C. Marticorena et al. 779
(CONC); Laguna Conguillio, 38º39’S, 71º38’W, 1000 m,
II-1963, Gleisner 201 (CONC); Prov. Malleco, Angol,
Lealtad, 6-XI-1958, Gunckel 374 (CONC); Angol, Deuco,
8-XII-1960, Montero 6396 (CONC); Angol, 80 m, 4-I-1923,
Barros 3370 (CONC); camino entre Lonquimay y cordillera
de las Raíces, 4-I-1947, Pfister s.n. (CONC 7201);
Lonquimay, 2-III-1939, Burkart 9491 (SI); camino entre
Lonquimay y Liucura km 31, 1000 m, 10-II-1960, Ricardi
& C. Marticorena 5052 (CONC); camino de Icalma a
Liucura, 16-I-1947, Pfister s.n. (CONC 7396); valle de
Lonquimay, 1000 m, 7-I-1947, Pfister s.n. (CONC 7273);
Malleco, Termas de Río Blanco, 1-I-1948, Pfister s.n.
(CONC 7868); Curacautin, Termas Río Blanco, 28-I-1938,
Montero 3667 (CONC 78523, 92599); a orillas del estero
Lancú, en la confluencia con el río Lolco, 950 m, 9-I-1977,
C. Marticorena et al. 1351 (CONC); Mininco, 20-I-1953,
190 m, Gunckel 819 (CONC); camino al Parque Nacional
Nahuelbuta, 1 km más arriba de los Lleulles, 37º46’S,
72º47’W, 800 m, 16-I-1986, R. Rodríguez & Pacheco 1912
(CONC). Región de los Lagos. Prov. Chiloé, Pudeto,
Purranque, 9-II-1961, C. Marticorena 1591 (CONC); Prov.
Llanquihue, Petrohué, Lago Todos Los Santos, 210 m, 24II-1944, Pfister 4277 (CONC). Región de Aysén del General
Carlos Ibáñez del Campo. Prov. Coyhaique, Coyhaique, 16I-1946, Barros 6076 (LP); S/ Prov., Valle Ibáñez, 740 m,
25-I-1939, Von Rentzell 6138 (SI 022629); Est. Guido, 17I-1952, Pfister & Ricardi s.n. (CONC 12181). Región de
Magallanes y Antártica Chilena. Sin loc. determinada,
1838-1842, Herbarium of the U.S. South Pacific Exploring
Expedition under the command of Capt. Wilkes, U. S. N.
(NY, PH 01084668).
NAMES OF DUBIOUS IDENTITY
Gnaphalium puberulum DC., Prodr. 6: 224. 1838. TYPES:
‛in Chili ad montem la Leona specim. anthesi incipiente
legit cl. Bertero (herb. n. 299!) et in Brasilia si Gn.
decurrens Schrank pl. Mont. t. 84 (non Yves) huc ex icone
ritè referendum?’.
According to the protologue two syntypes were cited, one
from Chile (Bertero 299) and other from ‛Brasilia’. We
found at P (P00704533, = phot!) the collection from Chile
‛In rupestribus herbidis Montis La Leona. Rancagua. Chili.
Bertero herb. N°299 / 1818, 8bre.’ (year 1833, in the sheet
at G-DC, not seen), which is in accordance in collector,
number and locality with the protologue but, as was pointed
out by Cabrera in this specimen, through an annotation on
the specimen’s sheet, its characters differ (probably refering
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
to the pubescence) from the protologue. It says ‛caule …
etomentosis pube brevi subglandulosa puberulo-hirtellis’
…. ‛Pubes fortè viscosa, sed á G. viscoso…’ vs. pubescence
long lanuginose (probably with short glandular trichomes
hidden under the wool) on stems and leaves in the material.
Gnaphalium puberulum DC. resembles Pseudognaphalium
viravira in having approximate basal leaves and remote
stem leaves that are apically attenuate, and capitula arranged
in dense terminal clusters. However since we were unable to
see the original material and we have seen no other material
which can be assigned to this species, the identity of G.
puberulum is not clear.
Gnaphalium chilense Spreng., Syst. Veg. 3: 480. 1826.
Gnaphalium sprengelii Hook. & Arn., Bot Beechey Voy.:
150. 1833, nom. superfl. TYPE. ‛Chili. Chamisso. Peru?’
(not seen).
Gnaphalium sprengelii Hook. et Arn. (Bot Beechey Voy.:
150. 1833) is a nomen supefluous because this name was
created to rename Sprengel’s Gnaphalium chilense (Syst.
Veg. 3: 480. 1826), since Hooker and Arnott considered that
the species was originally native from California and not
from Chile. Unfortunately, type collection of Gnaphalium
chilense Spreng., ‛Chili. Chamisso. Peru?’, was not located
at P, where the original herbarium of Sprengel is probably
deposited. For this reason, until the type of Sprengel’s name
is located or any materials can be provided, the identity of
G. chilense is uncertain.
EXCLUDED SPECIES
Gnaphalium oliganthum Phil., Linnaea 33: 167. 1864.
TYPE: ‛In collibus aridis prope Catemu prov.
Aconcagua legi.’ = Gamochaeta oligantha (Phil.) L.E.
Navas (Freire & Iharlegui 2008).
Gnaphalium pedunculatum (Poepp. et Endl.) Benth.
et Hook. ex Klatt, Linnaea 42: 142. 1878. Lucilia
pedunculata Poepp. et Endl., Nov. gen et sp. 3: 51. 1832.
Type: CHILE. Andes de San Jago, Jan 1829, Poeppig
s.n. (W not seen, digital image!, photo SI!) = Chevreulia
pusilla DC.
OBSERVATION
According to the protologue and digital image of Lucilia
pedunculata it corresponds to Chevreulia pusilla by having
capitula longly pedunculate.
Gnaphalium ramosum Phil., Linnaea 33: 164. 1864, nom.
illeg., non Lam. 1779 nec Sch. Bip. 1845. TYPE: Chile.
Aconcagua. Concumen, I-1863, C.L. Landbeck s.n.
(lectotype, here designated, SGO 64485!; isolectotype
LP 001942!) = Gamochaeta
OBSERVATIONS
After examining the type material, we propose to exclude
Gnaphalium ramosum from Gnaphalium since it lacks the
diagnostic character of the genus in having bristles pappus
fused at the base instead of bristles pappus with bases
cohering by patent cilia.
Muñoz Pizarro (1960: 143) mentioned ‘Hay un ejemplar
colectado cerca de Concumén, en la provincia de Aconcagua
(67787)’ of Gnaphalium ramosum Phil.We selected as
lectotype SGO 64485 since the sheet SGO 67787 was not
located.
Gnaphalium suffruticosum Phil., Linnaea 33: 165.
1864. TYPES: ‛Prope Illapel et Choapa invenit orn.
Landbeck’ = Gamochaeta suffruticosa (Phil.) Anderb.
(Freire & Iharlegui 2008).
ACKNOWLEDGEMENTS
We thank the reviewers for useful comments on a first draft
of the manuscript. Appreciation is expressed to the directors
and curators of the herbaria for the loan of specimens and
copies of the types that made this study possible. Special
thanks are due to Werner Greuter (B) who kindly helped
us with nomenclatural aspects of Pseudognaphalium
cabrerae. We would like to express our sincere appreciation
to Matthias Svojtka (W), for providing literature not
available in our countries. We thank María Alejandra
Migoya for the illustrations of Pseudognaphalium
aldunateoides, P. cabrerae, P. cymatoides, P. gayanum,
P. landbeckii, P. remyanum, P. viravira, and drawing
the maps. Financial support (PIP 112-200801-02196)
was provided by Consejo Nacional de Investigaciones
Científicas y Técnicas (CONICET), Argentina, the Chilean
Comisión Nacional de Ciencia y Tecnología (CONICYT:
Fondecyt 111085016/120448), and Universidad Nacional
de La Plata, Programa de Incentivos, Secretaría de Políticas
Universitarias, Ministerio de Educación, Argentina.
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INDEX OF SCIENTIFIC NAMES
Synonyms are in italics, accepted names in bold face.
Achyrocline 72
Anaphalis DC. 69
Chevreulia pusilla DC. 103
Gamochaeta Wedd. 69, 72
Gamochaeta oligantha (Phil.) L.E.Navas 103
Gnaphalium L. p.p. excl. type 69, 72
subg. Laphangium (Hilliard & B.L. Burtt) P.D. Sell 72
sect. Calolepis Kirp. 72
acutifolium Phil. 76
aldunateoides J. Rémy 73, 75
andicola Phil. 98, 100
araucanum Phil. 76, 79
argyrolepis Phil. 87, 89
cabrerae S.E. Freire 76
canum Phil. 81, 83
cheiranthifolium Lam. 76, 79, 100
var. multiflorum J. Koster 79
var. paniculatum (Bertero ex Colla) Skottsb. 79
chilense Spreng. 103
citrinum Hook. & Arn. 76
coquimbense Phil. 98, 100
cymatoides Kunze ex DC. 81
var. glabrum Walp. 81
diminutivum Phil. 75
dysodes Spreng. 94
fastigiatum Phil. 76
gayanum J. Rémy 83, 85
glandulosum Klatt 91, 94
heterophyllum Phil. 85
heterotrichum Phil. 72, 85
illapelinum Phil. 98, 100
insulare Phil. 73, 75
lacteum Meyen & Walp. 87, 89
landbeckii Phil. 89
leucocephalum Phil. 85
longifolium Phil. 98, 100
luteoalbum L. 73
moelleri Phil. 81, 83
montevidense Spreng. 98, 100
oliganthum Phil. 103
oxyphyllum DC. 69, 72
paniculatum Colla 79
pedunculatum (Poepp. & Endl.) Benth. & Hook. ex Klatt 103
perpusillum Phil. 73, 75
phaeolepis Phil. 75
philippii Cabrera 76
pratense Phil. 98, 100
pseudohelichrysum Reiche 85
psilophyllum Meyen & Walp. 91, 94
puberulum DC. 103
ramosum Phil. 103
remyanum Phil. 94
rivulare Phil. 75
robustum Phil. 85
sprengelii Hook. & Arn. 103
subnudum Phil. 98, 100
suffruticosum Phil. 103
tarapacanum Phil. 96
ulophyllum Hook. & Arn. 81
valdivianum Phil. 79
viravira Molina 98, 100
Helichrysum Mill. 69, 72
Hypelichrysum Kirp. 72
heterotrichum (Phil.) Kirp. 72
Laphangium (Hilliard & B.L. Burtt) Tzvelev 69, 72
Lucilia pedunculata Poepp. & Endl. 103
Pseudognaphalium Kirp. 69, 72
subgen. Laphangium Hilliard & B.L. Burtt 72
aldunateoides (J. Rémy) C. Monti N. Bayón & S.E. Freire
69, 70, 71, 72, 73, 74, 75
105
Gayana Bot. 71(1), 2014
andicola (Phil.) C. Monti N. Bayón & S.E. Freire 98
cabrerae (S.E. Freire) Deble 69, 70, 71, 73, 76, 77, 79
cheiranthifolium (Lam.) Hilliard & B.L. Burtt 69, 70, 71,
72, 73, 76, 78, 79, 100
coquimbense (Phil.) Anderb. 98
cymatoides (Kunze ex DC.) Anderb. 69, 70, 71, 73, 81, 82,
83, 85
fastigiatum N. Bayón 76
gayanum (J. Rémy) Anderb. 69, 70, 71, 73, 81, 83, 84, 85
glandulosum (Klatt) Anderb. 91
heterotrichum (Phil.) Anderb. 72, 85
illapelinum (Phil.) Anderb. 98
lacteum (Meyen & Walp.) Anderb. 69, 70, 71, 73, 87, 88, 89
landbeckii (Phil.) Anderb. 69, 70, 71, 73, 89, 90
luteoalbum (L.) Hilliard & B.L. Burtt. 73, 98, 100
moelleri (Phil.) Anderb. 81
montevidense (Spreng.) Anderb. 98
munoziae N. Bayón C. Monti & S.E. Freire 70, 71, 72, 73,
91, 92
oxyphyllum (DC.) Kirp. 72
perpusillum (Phil.) C. Monti N. Bayón & S.E. Freire 73
pratense (Phil.) Anderb. 98
psilophyllum (Meyen & Walp.) Anderb. 70, 71, 73, 91, 93, 96
puberulum DC. 102
remyanum (Phil.) Anderb. 69, 70, 71, 73, 94, 95, 96
robustum (Phil) Anderb. 85
tarapacanum (Phil.) Anderb. 70, 71, 73, 96, 97
viravira (Molina) Anderb. 69, 70, 71, 72, 73, 98, 99, 100, 103
INDEX TO NUMBERED COLLECTIONS EXAMINED
The number in parentheses refers to the corresponding species number in text.
A.J.H. 680 (12). Aedo 6806 (5); 7351 (3). Angulo 105 (3). Antoniz
s.n. (6). Aravena 271 (3). Arellano 48720 (12). Arriaga s.n. (5).
Arriagada s.n. (4). Arroyo 84-691 (6); 85-586 (9); 81112-A (11).
Arroyo & Humaña 99887 (4); 991714 (1). Arroyo et al. 85-393
(9); 97069 (9). Atanasio 706 (2).
Baeza & López 2759 (3). Baeza et al. 3074 (12); 3110 (12). Barnier
225 (3); 427 (12); 488 (7). Barrientos 1621 (4); 1699 (4); 2025 (4).
Barros 12 (12); 18 (12); 34 (1); 40 (5); 44 (12); 45 (1); 184 (12);
195 (5); 477 (12); 649 (1); 1466 (12); 1467 (5); 1940 (2); 2255 (4);
2258 (1); 2260 (4); 2272 (1); 2279 (3); 2285 (12); 2290 (1); 2301
(12); 2303 (9); 2304 (12); 2307 (4); 2318 (5); 2321 (5); 2417 (12);
2424 (12); 2840 (12); 3139 (12); 3370 (12); 3404 (5); 3874 (5);
6076 (12); 7332 (9); 21755 (12); 22611 (12); 24725 (5). Bayer 42
(5). Behn in 1941 (3); in 1928, in 1945 (4); in 1948 (5); in 1943.
(12); 20879 (12); 24413 (5). Belisario 849 (12). Belmonte 20060
(9); 20154 (5). Bertero 1823 (12, type of Gnaphalium viravira).
Bliss 674 (4); 2272 (1). Bliss & Lusk 575 (4). Böcher et al. 543 (5).
Boelcke 363 (12); 2424 (1); 3846 (5); 3885 (5). Buchtien s.n. (5);
3 (5), 174 (10). Bultmann 23661 (5). Burkart 9383 (4); 9491 (12).
Cabrera 3506 (5); 3636 (12); 11415 (5); 11462 (7); 12215 (12);
12523 (12); 12528 (5); 12697 (5); 19677 (12). Calderón s.n. (5).
Cañulaf s.n. (4). Chang 20965 (12). Collantes s.n. (5). Crisci 349
(3); 490 (5). Cuming 334 (4); s.n. (1).
coquimbense); 681 (5, type of Gnaphalium heterotrichum); 739
(1, type of Gnaphalium perpusillum). Geisze s.n. (5, type of
Gnaphalium heterophyllum). Germain s.n. (1, type of Gnaphalium
insulare). Gleisner s.n. (12); 201 (12). Grandjot s.n. (12); 995 (5).
C. & G. Grandjot s.n. (12). Gunckel in 1924, in 1933 (3); in 1943
(4); in 1955 (5); 374 (12); 819 (12); 1488 (3); 2999 (3); 3196 (3);
3324 (3); 11011 (4); 12826 (3); 12827 (4); 19193 (3); 21612 (4);
22280 (12); 22760 (12); 23443 (3); 23453 (12); 28012 (3); 30004
(5); 32025 (5); 36408 (3); 39466 (12); 40170 (4); 40215 (3); 40589
(3); 40598 (12); 41862 (3); 43572 (3); 46477 (7); 46993 (1); 48696
(9).
Harvey s.n. (4). Hastings 418 (12). Hicken 122 (5). Hochleitner s.n.
(3). Hollermayer s.n. (4). Humaña et al. 20102 (12). Hutchinson 67
(5).
Ibáñez et al. s.n. (5). Igaymán & M. Muñoz s.n. (5).
Jaffuel s.n. (4); 896 (12); 899 (12); 1306 (3); 2681 (5); 3267 (5).
Jaffuel & Pirion 3026 (4); 3226 (4). Jiles s.n. (5); 542-a (5); 910
(12); 1112 (4); 1686 (4); 1818 (12); 1848 (5); 3047 (5); 3429 (5);
3887 (9); 4428 (12); 4735 (12); 6083 (6); 6437 (5). Joseph 3657
(5). Junge in 1934 (3); in 1940 (4); in 1936 (12); 6285 (12); 6767
(1).
Kausel 3806 (5). Kuntze s.n. (12).
De Barba 2040 (12). De Vore 1545 (5). Dessauer s.n. (5). Díaz s.n.
(1, type of Gnaphalium diminutivum).
Elgueta 546 (2). Escudero s.n. (5).
Fernández 1494 (5). Finckh 552 (12). Finot & Leppe 97 (3); Finot
& López 1804 (12); s.n. (12). Frumau 63 (1, type of Gnaphalium
phaeolepis). Fuhrmann s.n. (3).
Garaventa 320 (5); 1151 (9, type of Gnaphalium psilophyllum);
1965 (3); 3117 (5); 5681 (12); 1293 (5). García 4083 (5). Gardner
et al. 295 (12). Gay s.n./s.f. (4); 678 (12, type of Gnaphalium
106
Lammers et al. 6381 (5). Landbeck in 1860 (5, type of Gnaphalium
leucocephalum); XII-1861 (7, type of Gnaphalium landbeckii);
XII-1862 (12, type of Gnaphalium illapelinum). Landero et al. 48
(6). Landrum & Landrum 8887 (9). Ledezma 8 (4); 225 (11). Levi
Heins 150 (4); 154 (3); 174 (4); 175 (5); 306 (11); 445 (5); 2911
(3). Looser 203 (12); 3765 (5); 3990 (12); 4394 (5); 66209 (3);
66226 (4); 66253 (5). López 11403 (3). Luebert & Teillier 2232
(12).
Mahu s.n. (12); 4885 (12); 8657 (5); 9795 (12); 10213 (12); 10335
(3). A. Marticorena & Jiménez s.n. (5); 214 (3). C. Marticorena
Revision of the genus Pseudognaphalium in Chile: FREIRE, S. ET AL.
1591 (12). C. Marticorena & Furet 62 (3). C. Marticorena &
Matthei 49 (9); 67 (9); 967 (12); 973 (12). C. Marticorena &
Rodríguez 8417 (12); 8430 (12). C. Marticorena et al. s.n. (12);
14 (6); 21 (9); 29 (11); 43 (12); 46 (5); 141 (9); 142 (6); 227 (6);
244 (6); 276 (4); 316 (6); 353 (3); 360 (11); 421 (12); 468 (5); 779
(12); 1006 (3); 1307 (3); 1338 (4); 1351 (12); 1612 (5); 9956 (5);
9972 (4); 83413 (12). C. Martínez s.n. (5). G. Martínez 49072 (5).
Matte 745 (12). Matthei & Quezada 463 (5); 1420 (1). Matthei
& Rodríguez 269 (4); 299 (3). Meyer 9684 (3). Mihoc 4 (4); 60
(4). Mihoc & Teneb 3897 (3). Mihoc et al. 3308 (3); 6227 (1).
Montero 222 (3); 1881 (5); 3667 (12); 5728 (1); 6056 (12); 6364
(4); 6396 (12); 9005 (3); 9437 (5); 10151 (3); 11575 (3); 12045
(2); 12281 (12). Moreira 1953 (8). Moreira et al. 1676 (8, type
of Pseudognaphalium munoziae). Moreno s.n. (5); 25689 (12). H.
Muñoz 48718 (12); Muñoz & Sierra 7327 (3).
Navarro s.n. (12, type of Gnaphalium andicola). Navas in 1953 (4);
in 1954 (4); 632 (4); 678 (11); 1389 (5). Niemeyer & Fernández
8908 (12). Niemeyer et al. 8970 (9); 89101 (6); 89106 (6); 89122
(5). Novoa s.n. (3); s.n. (12).
Oehrens s.n. (12). Ojeda 20949 (7).
Palma & Inostroza s.n. (3); Parra 37 (1); 475 (5). Parra & Torres
163 (3); 223 (3); 431 (12); 782 (3); 819 (3); 1026 (3). Pennell 12845
(4). Pfanzelt 173 (12). Pfister in 1935, in 1946. (3) in 1951, in
1955, in 1956 (4); in 1948, in 1952 (5); in 1950 (9); in I-1950 (11);
in 1944, in 1947, in 1948, in 1949 (12); 786 (3); 4277 (12); 4278
(3); 10502 (1). Pfister & Ricardi s.n. (3); s.n. (12). Philippi 18381842 (4); I-1860 (12, type of Gnaphalium pratense); in 1877 (10,
type of Gnaphalium remyanum); III-1878 (2, type of Gnaphalium
fastigiatum); II-1882 (12, type of Gnaphalium longifolium); I-1884
(12); II-1885 (11, type of Gnaphalium tarapacanum); III-1885 (4,
type of Gnaphalium moelleri); 17-II-1885 (6); III-1887 (4, type of
Gnaphalium canum); I-1888 (5, type of Gnaphalium robustum);
I-1893 (3, type of Gnaphalium acutifolium); s.n./s.f. (5). Pisano et
al. 1567 (12).
Rabanales s.n. (12). Rahmer III-1885 (6, type of Gnaphalium
argyrolepis); I-1886 (11, type of Gnaphalium tarapacanum)
Ricardi 20-XII-1950 (3); 29-XII-1950 (12); 2448 (12); 2853 (5);
3157 (12); 5445 (1); 9259 (1). Ricardi & C. Marticorena 3747
(12); 3956 (5); 4222 (12); 4223 (5); 4488 (5); 4730 (9); 4807 (6);
4943 (4); 4943 bis (4); 4972 (5); 5052 (12); 5150 (12); 5721 (12);
5766 (12). Ricardi & Silva 3535 (9); 3544 (6). Ricardi et al. 155
(5); 279 (11); 286 (9); 325 (6); 398 (9); 788 (6); 790 (5); 796 (5);
834 (12); 1348 (12); 1551 (12). Richter s.n. (12). R. Rodríguez
661 (12); 1476 (3); 2789 (5). R. Rodríguez & Baeza 2371 (4).
Rodríguez & C. Marticorena 3053 (12). R. Rodríguez & Pacheco
1912 (12). Roesner s.n. (5); 6508 (1). Rosas 1385 (5); 1392 (5);
1465 (5); 1507 (5); 4799 (5); 4946 (11). E. Ruiz & P. López 1018
(12); 1184 (12); 1231 (11).
Saa s.n. (5). Saavedra & Pauchard 111 (12). Santos 154 (3).
Schlegel 433 (5); 1537 (3); 3125 (12); 3144 (12); 3199 (4); 3329
(5); 4895 (4); 4949 (12); 5863 (12); 8034 (5). Semler s.n. (9). J. &
A. Solomon 4226 (3). Sparre 121 (3); 11016 (5). Sparre & Smith
80 (3). Stuessy & Crawford 6436 (3). Stuessy & López 11449 (3);
Stuessy et al. 6201 (3). Swenson & Stuessy 399 (3). S/leg. (4).
Teillier 3224 (1); 4624 (5); 5111 (5); 5851 (5); 5915 (9); 6489 (5);
51112 (1). Teillier & González 2301 (12). Teillier & Márquez 4848
(5); 4870 (4); 5313 (5); 5314 (5); 5315 (12); 5316 (1). Teillier &
Niemeyer 3322 (4). Teillier et al. 2027 (5). Teneb 551 (12); 569
(12). Tepe 2003 (12). Tepe et al. 1688 (5); 1689 (12); 1727 (3);
1840 (12); 1923 (3). Tomé 128 (5); 129 (5); 130 (12).
Ugarte 174 (3). Urtubey et al. 700 (1).
Vera & Ortiz 21 (3); 136 (3); 151 (3). Villagrán 5699 (3). Villagrán
& Armesto 2271 (9). Villagrán & Leiva 7269 (3); 7448 (9); 7545
(3). Villagrán et al. 1271 (11); 2353 (9); 2441 (9); 2478 (9); 9098
(9); 9229 (8); 9237 (9). Villarroel & Weldt 123 (3). Volkmann s.n.
(3, type of Gnaphalium araucanum). Von Bayern 388 (5). Von
Bohlen 742 (5). Von Rentzell 6138 (12).
Wagenknecht s.n. (3); 4396 (5). Weisser Sievers s.n. (5). Weldt &
Rodríguez 946 (3); 1149 (11). Werdermann 37 (3); 107 (5); 470
(5); 1092 (6); 1100 (9). Wilkes s.n. (12). Worth & Morrison 16494
(5).
Zemelman s.n. (4). Zollitsch 22 (5). Zöllner 976 (9); 978 (5); 1068
(4); 1069 (5); 1076 (5); 1138 (5); 1770 (12); 1772 (5); 2200 (5);
3031 (12); 5133 (12); 7936 (12); 8088 (5); 9914 (12); 10151 (5);
10198 (5); 10442 (5); 11254 (9); 11790 (5); 11991 (9); 12142 (12);
12518 (5); 13111 (12); 13147 (12); 13652 (12).
Recibido: 31.07.13
Aceptado: 28.04.14
107